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ZOOLOGICAL EESULTS
BASED ON MATERIAL COLLECTED IN
NEW BEITAIN, NEW GUINEA, LOYALTY ISLANDS
AND ELSEWHERE.
PART IV.
ILoniion: C. J. CLAY AND SONS,
CAMBRIDGE UNIVERSITY PRESS WAREHOUSE,
AVE MAEIA LANE.
AND
H. K. LEWIS,
136, GOWER STEEET. W.C.
©lasgoto: 50, WELLINGTON STEEET.
ILfipjig: F. A. BROCKHAUS.
p,fto Porfe: THE MACMILLAN COMPANY.
aomhap: E. SEYMOUE HALE.
ZOOLOGICAL EESULTS
BASED ON MATERIAL FROM
NEW BRITAIN, NEW GUINEA, LOYALTY
ISLANDS AND ELSEWHERE,
COLLECTED
DURING THE YEARS 1895, 1896 AND 1897,
BY
ARTHUE WILLEY, D.Sc. Lond., Hon. M.A. Cantab.
LECTURER ON BIOLOGY IN GUY's HOSPITAL, LONDON.
A
PART IV.-Vl,
{MAV. moo.)
CAMBRIDGE: ^X o, A^
AT THE UNIVERSITY PRESS. "^
inoo
CAMBBIDCtE :
PRINTED BY J. AND C. F. CLAY,
AT THE UNIVERSITY PRESS.
3
ph. 4-^
/J/
CONTENTS OF PAET IV.
PAGE
18. On the anatomy of a supposed new species of Coenopsainmia
from Lifu .......... 357
By J. STANLEY GARDINER, M.A.
With Plate XXXIV. and two figures in the test.
19. On the Insects from New Britain . . . . . . 381
By D. SHARP, M.A., M.B., F.R.S.
With Plate XXXV.
20. On the Stomatopoda and Macrura brought T>y Dr Willey from
the South Seas ......... 395
By L. A. BOBRADAILE, M.A.
With Plates XXXVI.— XXXIX.
21. Report on the Slugs' . ., 429
By WALTER E. COLLIXGE, F.Z.S.
With Plate.s XL.— XLI.
22. Report on the Polyzoa collected by Dr Willey from the
Loyalty Isles, New Guinea and New Britain . . . 439
By E. G. PHILIPPS.
With Plate.s XLIL— XLIII.
23. The Hydroid Zoophytes collected hy Dr Willey in the
Southern Seas ......... 451
By LAURA ROSCOE THORXELV.
With Plate XLIV.
24. Astrosclera willeyana, the type of a new Family of Sponges . 459
By J. J. LISTER, M.A., F.Z.S.
With Plates XLV.— XLVIII. and three figuie.'i in the text.
' I am desired to state that this article was written iu November 1m98 and received by me from the
author upwards of twelve months ago. A. W.
/■^ Ypi CONTENTS.
PAGE
25. A contribution towards our knowledge of the pterylography
of the Megapodii 483
By W. P. PYCRAFT, A.L.S.
With Plate XLIX.
26. The Stolonifera and Alcyonacea collected by Dr Willey in
New Britain, etc 493
By SYDNEY J. HICKSON, M.A., D.Sc, F.R.S.,
and
ISA L. HILES, B.Sc.
With Plates L.— LI.
27. Report on the Xeniidae collected by Dr Willey . . . 509
By J. H. ASHWORTH, D.Sc.
With Plates LIT. and LIII.
ON THE ANATOMY OF A SUPPOSED NEW SPECIES OF
COENOPSAMMIA FROM LIFU.
By J. STANLEY GARDINER, M.A.,
Felloiu of Gonville and Cains College, Cambridge.
With Plate XXXIV.
IXDEX OF CONTEXTS.
PAGE
Preface 357
Section I. General Anatomy of the Skeleton and Specific Description 358
General form — Mode of growth — Peritheca — Costae — Theca — Cahce— Septa — Cohimella — ElJect
of boring organisms.
Sectiox II. General Anatomy of the Polyps 360
Composition of the colony — Peritheca and Coenosarc — The expanded polyji — The contracted
polyp (circular sphincter muscle) — Coenosarcal canals and mode of budding — Tentacles —
Retractor muscles — Stomodoeum — Mesenteries — Generative organs.
Section III. Minute Anatomy 367
Ectoderm (external, peristomial and tentacular) — Tentacular nematocysts (anatomy and de-
velopment)— Stomodoeinn — Mesenterial filaments — Mesenterial nematocysts (anatomy and
development) — Calicoblastic ectoderm and attachment of the structui'eless membrane to
the corallum — Endoderm (longitudinal and circular sphincter muscles) — Generative organs.
Section IV. Conclusions relating to the body Layers in the Actinozoa 374
PREFACE.
Amongst the material very generously handed over to me by Dr Willey for
examination were nine colonies, with from two to twelve polyps, of a species of Coeno-
2)sammia, which I believe to be new, and for which I propose the specific name of
C. willeyi. All were obtained at Sandal Bay, Lifii, from the surfaces of reef-patches
on the under-side of coral masses in company with Distichopora and Stylaster. The
colour of the living colonies was bright red, with orange mouth-discs or peristomes.
w. IV. 49
358 ox THE ANATOMY OF A SUPPOSED NEW SPECIES
All were preserved by dropping into 90 per cent, alcohol — sometimes with addition of
formalin — which seems in most cases to have penetrated rapidl}', so that they are
well preserved for histological purposes.
The genus Coenopsammia was first defined by Milne Edwards and Haime (6),
who described nine species and placed the genus in the family Eupsainmidae. The
classification of these authors was in the main retained by Martin Duncan (5), who
gi'ouped in a somewhat ai'bitrary manner their several families into various alliances,
and added the definitions of the numerous new genera described since the publication
of the Histoire des Coralliaires (6).
The genus Coenopsammia is one of the simplest members of the family Eupsam-
midae, and the sjiecies here described has no zoosanthellae in its endoderm and must
hence feed entirely by means of the food taken in through its stomodoeum. Further-
more it produces buds from the basal edge of the polyp as do the most primitive
colonial Actiniae, and might therefore reasonably be expected to retain much of the
structure of the Hexactinian pol^'ps from which all the Madreporaria seem pi'imitively
to have been derived.
For the so-called mesogloea or jelly I prefer to use the term structureless membrane
or basement membrane, as the layer apjjears to me to be of the same nature as basement
membranes in general.
SECTION I.
General Anatomy of the Skeleton and Specific Description. (PI. XXXIV., Figs.
1 — 3, and text-figures, I. II.)
The corallum' is devoid of any epitheca, the whole except the attached base being
covered by the polyps. It occurs in its younger stages in the form of a single
corallite, which is gradually built up by the polyp, increasing both in diameter and
height. At the same time the theca is thickened near the base of the corallite by
deposit from the extrathecal portions of the polyp ; and irregularly arranged intrathecal
platforms, or pseudotabulae, are formed across the calice.
Budding takes place near the base of the parent corallite, two, three or more
daughter corallites being constantly found of about the same size and age. The buds
at first project almost at right angles from the sides of the original corallite, but by
a more rapid growth of their outer or distal sides gradually turn upwards, yet always
at some slight angle to the parent corallite (Fig. 2). The greatest diameter of one such
corallite w^as 7'5 mm., and of its three buds from .3"5 to 5"o mm. The original corallite
was 14 mm. high, and the calices of the budded corallites formed a ring about 5 mm.
below the margin of its theca.
' For definition of terms relating to the skeleton see Martin Duncan (5).
OF COENOPSAMMIA FROM LIFU. 359
With the production of buds the base of the colony broadens. The parent and
daughter corallites continue to increase in height and size, daughter polyps being again
formed on their outer sides by budding from their free basal edge. The final result
is an incrusting mass with a number of corallites standing up separately upon it, the
oldest being tj'pically in the centre (Fig. 1).
As the corallites ai'e built up, skeleton of a loose, porous nature — peritheca' — is
deposited between them by the extrathecal portions of their polyps. The different
corallites of a colony are hence only free for a limited height, the highest free portion
of any corallite in the collection being 12 mm., and the greatest diameter of the largest
corallite 9 mm.
Hence, if growth proceeds regularly, a low convex mass is formed, the corallites
gradually decreasing in height and diameter fr(5m the centre outwards. The colonies
in the collection have this general form, but are all very small — largest 5.5 cm.
across (Fig. 1) — and rather irregular, being overgrown in places by foraminifera, sponges,
and other organisms. To the struggle between these and the polyps may be directly
ascribed in many places the variation in height of the corallites above the colony.
The mode of growth however of the species can be distinguished in all.
The corallites have on the outside an appearance of longitudinal striae, due to
the presence of low, rough, subequal costae, which correspond in number and position
to primary, secondary, tertiary, and quaternary septa. Many of the costae are con-
tinuous from the parent to the daughter corallites in the young stage, but where
much peritheca has been formed there is a distinct narrow valley between the corallites,
from which the costae diverge.
The theca is thin, and for a few millimetres below its upper edge very freely
perforated in lines between the costae (Fig. I.). It does not appear to be a true theca,
formed in the first place by the basal ectoderm as a ring on the basal plate, joining the
septa, but rather a pseudotheca, formed by the fusion of thickenings of the septal sides.
The calice is slightly oval in shape, the two diameters being in the proportion
to one another of nine to eight. Within it septa of three cycles are present, of which
the primaries and secondaries fuse with the columella (Fig. 3). The primary septa
generally project from the edge of the calice almost horizontally inwards for about a
quarter of its diameter — they often in the younger corallites rise slightly above the level
of the theca— ending by the axial fossa with almost smooth vertical edges. The two
primary septa, which lie between the directive mesenteries at each end of the longer
diameter of the calice — hence termed directive septa — do not project for more than
about one-seventh the diameter, so that the axial fossa is very distinctly oval''.
The secondary septa project horizontally from the upper edge of the calice for
about one-twelfth its diameter, ending then with almost vertical edges, but abruptly
broadening to join the columella. The tertiary septa are small and inconspicuous, and
the quaternary are low ridges, only seen in ground down surfaces or sections towards
the base of the calice.
' For definition of this term see p. 301.
« These proportions are not clearly shown in the figure, which has been somewliat diagrammatioally drawn
by the artist.
49—2
360 ON THE ANATOMY OF A SUPPOSED NEW SPECIES
The septa naturally decrease in thickness from the primaries to the tertiaries.
All are relatively thin and little perforated, with almost smooth edges and sides
covered with low, blunt, somewhat distant granules, not arranged in any determinate
manner. Synapticula are absent.
The columella closes in the axial fossa below, and owing to the narrow directive
septa, added to the slightly elongate shape of the calice, is very distinctly oval. It
varies in dejjth, in the larger corallites being situated from 4 to 5 mm. below the
edge of the calice. In the 3'oimgest separate corallite, that I have examined, there
appeared to be a true columella, arising from a basal plate, but in the older calices
it has a spongy appearance, and seems to have been formed principally by the ana-
stomosis of a large number of trabeculae from the septal edges.
I have not attempted to examine the minute anatomy of the corallum in any
detail. The skeleton in all the specimens is everywhere much bored into by algal
filaments, which although found principally in the deeper lying parts extend in places
to within '2 mm. of its surface.
SECTION II.
General Anatomy of the Polyps.
Composition of the Colony. The corallum, as mentioned before, is everj-where,
except over its attached base, covered by the polyps. The latter may be regarded
each as an independent individual capable of leading an independent existence. All
the polyps however are connected together by the coenosarc, which consists of a
number of canals separated from one another by a double layer of endoderm with
the structureless membrane between. These canals run from poh^) to pol3-p — branching
perhaps at the bases of the projecting corallites — and serve to put the gastrovascular
cavities of the diflerent pol}'ps in free communication with one another.
A similar arrangement is found in Pocillopora and all corals, so far as I am
aware, which have a well-developed peritheca, save that in some genera, as pointed
out by Fowler (10), the canals have been pushed apart from one another and the
external wall, consisting now of a double layer of ectoderm with the structureless
membrane between, lies directly on the corallum. In imperforate Madreporaria the
coenosarcal canals may be said to commence from the edge of the calice, while in
perforate forms, especially in such forms with partially free corallites as Coenopsammia,
no such sharp line of distinction can be drawn, since the iutracalicular portions of the
OF COEXOPSAMMIA FROM LIFV. 361
coelentera at frequent intervals communicate with the coenosarcal canals by ramifying
canals through the theca.
The coenosarcal canals in fact are simply extrathecal portions of the coelentera of
the diflferent poh-ps, which serve to connect theii- intrathecal or gastrovascular portions.
Peritheca and coenosarc. The corallum except over the base of attachment
is everywhere covered by the calicoblast layer of ectoderm. This is constantly depositing
skeleton over all parts more or less rapidly. Skeleton so deposited has been termed
by Martin Duncan (5), when it occurs outside the theca and between the costae, the
" exotheca," and, when it serves to fill up the valleys between the free portions of the
corallites, the " coenenchyma." Indeed, when the deposit of corallum outside the corallites
was small, Martin Duncan called it "exotheca," but, if considerable, " coenench}"ma."
In the asexual method of reproduction, which forms the colony, whether by fission
or bud formation, there is at first no coenenchyma between the two individuals, or
between the bud and its parent corallite. In individual specimens of any species the
coenenchyma varies enormously with the rate and form of growth of the colonj' ; it
hence seems to me improbable that its relative abundance alone can be in any genus
a specific distinction. There is no sharp line of sepai-ation nor of structure between
the "exotheca" and the "coenenchyma" in Coenopsammia, nor indeed in most Madre-
poraria, the latter as it is built up being necessarily fused with the former. In
Galaxea however the distinction is well marked, the "coenenchyma" having a porous
and the "exotheca" a compact structure.
It hence appears to me advisable that the term "coenenchyma," if retained in
the Madreporaria, should be applied either to the structure usually so-called in Galaxea
or to all parts of the corallum outside the theca. The term is of such wide application
that it would onlj" increase the confusion, which already prevails, to restrict it in such
a way. Either use of the term too is du-ectly opposed to its w-ell-established use
in the rest of the Anthozoa. I accordingly propose to use the term peritheca, which
was employed in the first place by Milne Edwards and Haime for the so-called
"coenenchyma" of Gala-xea. The peritheca is that part of the corallum of colonial
Madreporaria, which is deposited outside and subsequentlt/ to the theca. The coenosarc is
that part of the polyps in a colony which lies outside but not above (i.e. in expanded
state) the thecae of the several corallites. The " Randplatte " of Heider and von Koch,
the "edge-zone" of Miss Ogilvie, is then that part of the coenosarc which lies over
the free portions of the corallites. The above use of the term coenosarc is more in
accordance with its physiological meaning in the Alcyouaria and Hydrozoa.
The expanded polyp. As all the polyps of the specimens, entrusted to me by
Dr Willey, are completely retracted, it is impossible to speak definitely of the conditions
found in the living polyps. However, from the appearance of the contracted muscles
and the accordingly much bent mesenteries, I am led to believe that the polyps
expand themselves to a height of at least 4 mm. above the top of the theca. The
tentacles then form three circles close to one another round the peristome, or mouth-
disc, the outer with twelve, and the two inner each with six tentacles, the bases of
the outer and the two inner circles alternating with one another.
362
ox THE AXATOMY OF A SUPPOSED NEW SPECIES
Figure I. Diagrammatical transverse section through a single completely retracted polyp
in six different planes shown approximately in Fig. II.
The section is that of a single lateral sextant (or system) — without directive mesenteries —
of the same polyp traced downwards in a series of transverse sections.
The sextant is taken from the middle of one primary septum to the middle of the
next. (The position of the sections will be readily understood by reference to Fig. II., but
the polyp, from which this figure was made, was not quite in the same state of retraction.)
The corallum is dotted and covered everywhere by the calicoblastic ectoderm, structureless
membrane or lamella, and endoderm. th. Theca. I, II and III. Primary, secondary and
tertiary septa. coL Columella (only seen in F). e. w. Body-wall (consisting of ectoderm,
structureless membrane and endoderm) external to the mouth-disc or peristome. Limited by the
bases of the tentacles, p. w. Body-wall of mouth-disc or peristome, st. iv. Body-wall of the
stomodoeum, much thickened owing to the very thick ectodermic epithelium. T. 1, T. 2,
and T. 3 Completely retracted primary, secondary and tertiary tentacles covered with batteries
OF COENOPSAMMIA FROM LIFU. 363
of nematocysts. 1, 2 and 3, Primary, secondary and tertiary mesenteries, consisting of two
layers of endoderm with the structureless membrane between. The retractor muscles are
indicated as fine lines at right angles to the mesenteries, -m.f. mesenterial filaments.
Sections.
A. Above the theca. The primary septa are represented although they do not, except
in young corallites, project above the theca.
B. About half-way between the top of the stomodoeum and the top of the polyp. The
theca is complete and divides the coelenteron into extra- and intrathecal portions, connected
together by numerous perforating canals, commonly running as represented. The extrathecal
portions of the coelenteron are divided into canals, which correspond in number and position
to the spaces between the mesenteries. The costae and secondary septum are well marked and
the tertiary retractor muscles have appeared.
C. Through the mouths of the retracted tertiary tentacles, which lie externally to those
of lower orders. The retractor muscles of the secondarj' mesenteries have appeared and the
tertiary septa are indicated. (The perforating canals of the corallum and the peritheca are
not represented in this and subsequent sections.)
D. Through the mouths of the retracted primary and secondary tentacles. The polyp
is cut across in two places owing to the somewhat raised lip round the stomodoeum.
E. Through the lower half of the stomodoeum. The lower free ends of the retracted
primary and tertiary tentacles are well marked. The tertiary mesenteries are free with
distinct filaments ; in the left pair ova are represented, covered by the much thickened
nutritive endoderm.
F. Towards the base of the polyp. The right half of the section is rather higher than
the left. In it the tertiary mesenteries are much narrower with smaller filaments and
without ova, which have now appeared in the secondary mesenteries. In the left half the
tertiary mesenteries have lost their filaments and muscles and will shortly disappear; the
secondary mesenteries are narrower and have lost their muscles, while in the primary
mesenteries ova have appeared.
The contracted polyp. In the completely contracted condition the opening of the
stomodoeum is situated about Vh mm. above the top of the columella. The peripheral
part of the mouth-disc is drawn inwards and downwards, causing a marked depression
round the stomodoeal opening. This condition is brought about by the presence of
a strong circular sphincter muscle — Rotteken's muscle of the Actiniaria — (Fig. II. m.s.)
together with that of sti'ong longitudinal muscles on the mesenteries, spreading out
under the oral disc and within the tentacles. The presence of a circular muscle in
the Madreporaria was first suggested by Moselcy (21) in Sphetiotrochus and subseciuently
described by Fowler in the same genus (9) and also in Dwncania (11). But, whereas
in these genera the sphincter opening leads into a cavity in which the tentacles
lie, the condition differs in Coenopsammia owing to the complete introversion of the
tentacles in a more irregular manner but similar to that of Seriatopora (9). In fact
364 ON THE ANATOMY OF A SUPPOSED NEW SPECIES
round the stomodoeal opening is a prominent ridge, in the depression round which
lie the openings of the introverted tentacles (Figs. i. and il.).
Coenosarcal canals and mode of budding. The mesenteries are 48 in numbei
and have a determinable position in respect to the 24 septa. There are 48 inter-
mesenterial spaces, 24 enfocoelic between mesenteries of the same pair, into which the
septa project, and 24 exocoelic spaces between mesenteries of neighbouring paLrs\ From
each intermesenterial space is given off a coenosarcal canal passing outside the theca.
The dividing walls of these are continuous with the mesenteries over the theca, and
indeed may be regarded as their perijjheral ends or extrathecal portions (Figs. I. and II.).
The coenosarcal canals end blindly at the free edges of the colony, but in the central
parts they put each intermesenterial space of each several polyp into communication
with at least one such space in a neighbouring polyp.
Budding takes place from the blind ends of the coenosarcal canals at the basal
margin of the single jwlyp or colony. A number of the coenosarcal canals fuse together
and a mouth breaks through. The intermesenterial spaces of the daughter polyp on the
inner side, i.e. towards the parent polyp, are each formed directly from a single
coenosarcal canal, while those at the sides are formed by the branching of these canals.
The mesenteries are formed from the dividing walls of the coenosarcal canals, i.e. from
the extrathecal portions of the mesenteries of the parent polyp. The young corallite
appears to be very rapidly formed, and has from the first a diameter of 2b mm.'
The intermesenterial spaces, besides being in communication w^th the coenosarcal
canals over the theca, have also connecting canals through the theca. The dividing-
walls of the coenosarcal canals outside the free portions of the corallites are attached
to the skeleton between the costae, which project into their lumina. The mesenteries
on the other hand have their broad bases attached to the septa close to their fusion
into the theca, and hence somewhat facing one another. The connecting canals near
the top of the theca from the exocoeles run straight through the theca near the
base of one of the bounding mesenteries and open into their corresponding coenosarcal
canals (Fig. I., b). In the entocoeles the arrangement is similar in the same position,
but the connecting canals often arise at a considerable distance up the sides of the
septa and perhaps run diagonally through them. This arrangement of connecting canals
near the top of the theca, joining the intermesenterial spaces with their corresponding
coenosarcal canals alone, strongly supports the view that in this species the theca is formed
simply by the fusion of thickenings on the sides of the septa. Lower down in the
polyps instead of separate connecting canals, a system of ramifying and anastomosing
canals is found, similar to that described by Fowler for Rhodopsammia (7), but not
so complicated, owing probably to the more delicate corallum.
1 There would thus be between neighbouring primary septa, in each system 8 mesenteries, 2 primary,
2 secondary, and 4 tertiary. In one poln^ however in one system tliere were 10 mesenteries owing to an
addition of 2 tertiary, hut the next system had ouly 6 mesenteries, 2 tertiary beiug absent.
- In the various colonies in the collection I have only found one bud without a corallite, and in this
the preservation was not sufficiently good for me to follow out the process in any detail. I have traced
however the connections of two buds, each of about 3 mm. in diameter, with the parent polyp.
OF COENOPSAMMIA FROM LIFU.
365
Tentacles. The tentacles are all entocoelic, and their three orders con-espond
to the orders of septa, the tertiaries being situated most externally on the mouth-
disc. The introverted tertiary tentacles form deep pockets in the entocoeles of the
Figure II. Diagrammatical longitudinal section of a polyp, the left half passing through
the opening of a tertiary tentacle and exposing the face of a tertiary mesentery, and tiie
right half passing to one side of a primary mesentery exposing it likewise.
A — F. Sections shown in Fig. I.
th. Theca. .S'. /. and S. III. Primary and tertiaiy septa, col. Columella, i^th. Peritheca.
p. Perforating canals of the theca. e. tu. External body-wall. y. Ova. ;;. w. Body-wall of moutli-
disc. St. Storaodoeum. T. 1 and T. .3. Primary and tertiary tentacles retracted, m. Jilesenterial
filaments, m. r. Retractor muscle. i». s. Circular sphincter muscle.
Left Half. The tertiary mesentery does not extend beyond the tertiary tentacle. Its
muscles are well marked and attached nearly along its whole length to the theca, arising
from the wall of the tentacle. Along the free edge a much convoluted filament is present
except at its upper and lower ends. The ridge of ci>ralluni, represented at its lower edge,
is not generally marked, and the mesentery does not reacii the base of the polyp. The
outline of a tertiary septum {S. Ill) is shown.
Right Half. The lip rounfl the storaodoeum is well marked with a circular depression
around, having tlie opening of a primary tentacle ( 7'. 1 ), the position of wliich is shown
W. IV. 50
366 ox THE ANATOMY OF A SUPPOSED NEW SPECIES
by a broken line. The outline of a primary septum (.S'. /) is also shown, fusing with the
columella below. The retractor muscles arise from the whole of the mouth-disc, but are
only attached to the lower two-thirds of the theca. The free edge of the mesentery is
everywhere covered by a mesenterial filament, which is directly continuous with the stomodoeal
epithelium above.
tertiary mesenteries both above and below their open mouths (Fig. i. c and Fig. ii.).
Shorter pockets are found too, especially in sections where the open mouth is cut
through, projecting into the exocoeles outside the tertiary entocoele. In fact the
tertiar}' mesenteries would appear to continue for some distance into the extended
tertiary tentacles.
The introversion of the secondary and primary tentacles gives rise to deep some-
what irregular invaginations — extending only below their open mouths — in the entocoeles
over their corresponding septa (Fig. I. D, and Fig. II.). The iuvaginated portions are
crescent-shaped in transverse sections owing to the projecting septa, and have no
side diverticula into the exocoeles. The mesenteries are attached, one at each side
of the bases of these tentacles, and cannot extend into them in any way, when they
are expanded. The tentacles appear as if they decrease considerably in length from
the tertiaries to the primaries.
All the tentacles are covered with round, knobbed batteries of nematocysts, which
gradually decrease in size from their tips. At their bases these batteries are very
small, and pass almost imperceptibly — especially on the tertiary tentacles — into the
general ectoderm.
Retractor muscles. The great longitudinal retractor muscles of the mesenteries
have the regular arrangement, typical of the Hexactiniae, i.e. on the sides facing one
another on all the pairs except the directives. Their course may be best seen by
reference to Figs. i. and II. The fibres of the tertiary mesenteries arise from the
tentacles alone, while those of the primaries and secondaries arise from the whole
of the mouth-disc. They then run on all the mesenteries diagonally across to end
near their attachments to the corallum. The lower part of all the mesenteries is free
from muscular fibres.
On the faces of the mesenteries, opposite to the great i-etractor muscles, there are
a few isolated longitudinal muscles with a similar course. They are rather more
numerous on the directive mesenteries, the great retractor bands of which are not as
large as those of the other primary mesenteries. There do not appear to be any
definite protractor or transverse muscles. The circular sphincter muscle — Rotteken's
muscle — is a broad, diffuse band situated immediately below and outside — morphologi-
cally speaking — the tertiary tentacles (Fig. ii.).
Stomodoeum. The stomodoeum is about 1 mm. in length in the contracted polyp.
Its mouth is an elongate or oval slit, lying in the long axis of the directive septa,
without any trace of gonidial grooves at its ends. Below the mouth its lumen is
very irregular, being often drawn out into deep pockets between the septa.
Mesenteries. The tertiary mesenteries are attached to the mouth-disc in the
tentacles, but do not reach to the aboral or basal body wall. Their free edges, except
OF COENOPSAMMIA FROM LIFU. 3G7
immediately at the top and bottom, end in much convoluted filaments. The primary
and secondary mesenteries are attached to the stomodoeum, with which their filaments
are continuous, and also reach to the basal wall of the polj'p. AH the mesenteries
decrease in breadth with the decrease of the muscular bands, and indeed the tertiary
mesenteries are lost with their disappearance (Figs. i. and II.). The filaments of the
primary and secondary mesenteries run straight into the polyp below the stomodoeum
for 1 — 1"5 mm. before the convolutions commence. The filaments are attached to the
mesenteries for their whole length, and are without free portions (acontia) at their
lower ends.
Generative organs. All the polyps, which I have examined by sections — eleven
from six colonies — are female, and have no trace of any male generative organs.
The ova in each mesentery, in which they are found, are arranged in a row in the
structureless membrane between the convoluted mesenterial filament and the great
retractor muscle (Fig. Ii.). In the youngest polyps examined (3 mm. across the calice)
ova only occurred on the primary mesenteries, but in all the others — eight — had passed
into the secondary mesenteries as well. In two large polyps, the one cut into longitudinal
and the other into transverse sections, I also found small ova in the structureless
membrane of some of the tertiary mesenteries. The ova do not seem to pass into the
directive mesenteries as soon as the other primaries, perhaps on account of their being
less broad through the elongation of the stomodoeum.
The ova are presumably formed as in the Hexactiniae from the endoderm, and
wander into the structureless membrane (13 and 14). They here wander into the
structureless membrane of any mesenteiy, provided that it is of sufficient breadth. I have
also found a similar arrangement in Prionastraea ahdita, and it seems very doubtful,
whether the order of the mesentery, in which the generative organs are found, is of any
importance.
SECTION III.
Minute Anatomy.
Ectoderm. (Figs. 4 and .5.) The external ectoderm, i.e. of the general surface
of the colonies outside the tentacles, is very well preserved, and seldom or never
detached from the underlying structureless membrane. It is about '03 mm. in thickness,
and has a very uniform and much vacuolated appearance. Cell outlines can seldom be
distinguished, but it appears to be an epithelium of a columnar facies with a distinct
layer of nuclei in the centre. The latter are small, oval, with a few granules and
deep staining nuclear membranes. A few smaller, round nuclei, staining homogeneously,
are also found, and perhaps belong to sense cells (Fig. 5). Small nematocysts are
scattered about, but occur principally opposite to the attachments of the dividing walls
of the coenosarcal canals, where the epithelium is rather thicker. They vary up to about
•02 mm. in length, and generally have the same structure as the nematocysts of the
tentacles. A few, however, are exactly similar, except in size, to the nematocysts found
on the mesenterial filaments.
50—2
368 ON THE ANATOIY OF A SUPPOSED NEW SPECIES
Gland cells of two kinds are found, mucous and granular. The former stain brown
in Heidenhain's iron haematoxylin and eosiu, and are fairly numerous all over the
coenosarc, yet gradually decreasing in number from the bases of the tentacles down-
wards. The granular gland cells correspond to the "kornige Drlisenzellen," described
by the Hertwigs in Actiniae (1-i). Their gi-anules are very small, and stain black in
iron haematoxylin and eosin (Fig. 4). In shape some are narrow and very elongate,
while others are almost round or sack-like. In one young polyp (about 4 mm. across
the calice) they are very numerous for about 3 mm. below the tentacles, but over
the coenosarc between the free portions of the polyps very scarce. In older polyps
they are commonly found only at the bases of the tentacles.
A few oval nuclei occur immediately above the structureless membrane, but there is
no well-marked punctate nervous layer except near the bases of the tentacles. Muscular
jDrocesses of the cells and distinct muscular fibres are completely absent. The surface
of the layer is sometimes covered with mucus, but generally it is sharply defined, and
there is no appearance of the ciliation described by the Hertwigs in the Actiniae.
The ectoderm of the mouth-disc is very similar to the external ectoderm, but rather
thicker — about 'O-i mm. — and with a distinct finely punctate layer of nerve fibrils,
immediately over the structureless membrane. Some of the cells too have distinct basal
muscular processes. Mucous gland cells are numerous, especially near the mouth of the
stomodoeum, but gi-anular gland cells are absent. The nematocysts are more numerous
than in the external ectoderm and of the same two kinds, the tentacular towards the
exterior and the mesenterial around the stomodoeal opening.
The ectoderm of the tentacles takes the form of batteries of nematocysts (Fig. 6).
Each battery is packed in the centre with nematocysts, close to the basal ends of which
are a number of rod-shaped, or oval nuclei. Below these and towards the sides of
the tentacles the nuclei are round or very slightly oval. All stain homogeneously with
iron haematoxylin, the oval, granular nuclei of the external ectoderm being very rare.
Under the centre of the battery on the structureless membrane is a thick, finely granular
mass with a few supporting fibrils, a great concentration of the nervous layer {n. I. Fig. 6).
A few mucous cells occur towards the sides of the batteries, but granular gland cells are
absent. The muscular processes of the cells form a well-marked, thin, deeply staining
layer on the structureless membrane. They appear to run mainly in a longitudinal
direction in respect to the expanded tentacles, and arise principally from the cells around
the central mass of nematocysts. In the contracted condition the central part of the
battery is sometimes slightly depressed, but generally the nematocyst mass is much
pushed out owing to the contractions of these muscles.
Tentacular nematocysts (Figs. 7 — 10). The tentacular nematocysts, when ripe,
with the thread well formed (Fig. 7) are very uniform iu size, about '027 mm. in
length by -003 mm. in diameter. The thread is spirally coiled round a central homo-
geneous mass of pi-otoplasm, and causes the external membrane of the nematocyst
to project over it. The number of coils varies from 20 to 30, but commonly there
are about 2-i. In most nematocysts the terminations of the thread cannot be distin-
guished, but in some the thread may be seen to end at the base of the capsule in
a very finely granular mass of protoplasm, while at the opposite end it runs inwards
OF COENOPSAMMIA FROM LIFU. 369
and slightly backwards at first, and then straight to the somewhat pointed and slightly
projecting free end of the nematocyst. Generally the basal end of the nematocyst
lies in a finely granular mass of protoplasm with a nucleus either oval or rod-shaped.
Usually a filament, comparable to the granular peduncle described by Lendenfeld in
Hydra (19), can be distinguished running down to the nervous layer; the structureless
peduncle of Hamann (12) cannot be seen, but numerous processes of the structureless
membrane pass everywhere into the ectoderm.
Although I have very carefully examined the uematocysts in the tentacles of
seven polyps, I have failed to find any of a structure differing from the above. The
whole nematocj-st is thrown out when the thread is extruded, and in thick sections
I have been able to follow every stage of the process. The end of the thread is
ejected first and then turn after turn of the spiral follows (Fig. 8). The thread itself
under ocular 4 and y'j oil immersion, appears for its whole length to be absolutely
homogeneous and structureless.
I have considered it necessai'y to describe the structure and the appearances
on extrusion in some detail, as Mobius (20) appears to have confused neraatocysts
similar to these in the tentacles of Caryopkyllia with those found on the mesenterial
filaments of the same form. He appears indeed to have considered them to be young
stages in the development of the mesenterial uematocysts. Bourne (4) too, following
Mobius, gave a figure of a similar nematocyst, apparently not quite ripe, as a developing
nematocyst of the mesenterial foi-m.
Although I have not been able to follow every stage in the process of formation
of these nematoc3-sts, yet it seems to be as follows. In the place of the ejected
nematocyst appears a homogeneous mass of protoplasm, extending almost from the surface
of the ectoderm to the structureless membrane, and having near its base a nucleus.
The central part of this acquires a definite membrane and subsequently forms the
nematocyst. No nucleus can in any stage be distinguished within this membrane,
but the basal nucleus is nearly always well marked. Its protoplasm next becomes
very finely granular (Fig. 9), and the fine granules then fuse together and aiTange
themselves in definite spiral lines close under the external membrane (Fig. 10). The
young nematocyst is now about one and a half times as long and thick as when ripe.
Lastly the gi-anules fuse, forming the thread, and the external membrane still further
contracts. The thread seems to be tightly coiled up under a very elastic extenial
membrane and to be extruded mainly by its own elasticity. Even in preserved speci-
mens, when the ripe uematocysts are broken or cut across, three or four coils ot the
thread will uncurl. The thread indeed resembles both in its development and appearance
the elastic fibres in the connective tissue of Vertebrates.
Stomodoeum (Fig. 11). The ectoderm of the stomodoeum is rather thicker than
that of the mouth-disc, which gradually merges into it. It is composed of an elongated
columnar epithelium with crowded, rod-shaped nuclei, all staining homogeneously. Over
the structureless membrane is a thick, finely granular nervous layer— not distinctly seen
in the figure— with a few round nuclei and supporting fibres. A few nematocysts like
those of the mesenterial filaments but much .smaller are fo\ind ; tentacular nematocysts
are extremely rare.
370 ON THE ANATOIY OF A SUPPOSED NEW SPECIES
Mucous cells are scarce, but the epithelium is in places crowded with goblet-like
vacuoles with numerous interstitial nuclei towards the exterior (Fig. 11). They are
commonly much swollen with some apparently fluid unstainable secretion, and no nuclei
especially belonging to them are visible. These secreting vacuoles are found principally
opposite to the attachments of the primary and secondary mesenteries. Indeed in some
polj'ps they form well marked lines over these mesenteries, extending up almost to the
mouth of the stomodoeum. In one specimen, in which a Copepod is lying partially
in the stomodoeum and partially in the coelenteron, the vacuoles are much smaller
and generally appear to have discharged their secretion.
Mesenterial filaments (Figs. 12 and 13). The mesenterial filaments of the
primary and secondary mesenteiies are the direct continuations of the lower edge of
the stomodoeum and have a very similar structure to that described by Bourne (3)
and Fowler (8) for other Madreporaria. They are of a somewhat crescentic shape in
transverse sections (Fig. 12) and their epithelium is sharply marked off fi-om that of
the mesentery below them. The structureless membrane of the mesenteries ends in a
T-shaped swelling with numerous fibres passing off into the filament from its ends.
The central part of the filament is crowded with goblet vacuoles and is the
" Driisenstreifen " of the Hertwigs (14) and other German authors. This gradually passes
at the sides into an elongated columnar epithelium, which bends round the structureless
membrane so that it covers the whole underside of the horizontal bar of the T. These
parts correspond to the lateral lobes or " Flimmerstreifen," described b}' the Hertwigs in
the Actiniaria and by E. B. Wilson (27) in the Alcyonaria. Mucous cells are more
abundant than in the stomodoeum and lie principally at the sides; small glandular
vacuoles densely crowded with relativeh' large granules are found also in the same
position, but vary very largely in abundance in different polyps. Nematocysts do not
become numerous until the convoluted portion of the filament commences, when the
goblet vacuoles gradually decrease and finally disappear (Fig. 13). Near the stomodoeal
end of the mesenteries the nematocysts lie almost entirely at the sides of the filament,
but towards the base they occur in dense masses thi-oughout.
The filaments of the tertiary mesenteries, as in Fungia, are very similar to those
of the primaries and secondaries but slightly smaller. The goblet vacuoles are not
nearly so numerous, and do not form a well marked clearer area in the centre.
Mesenterial Nematocysts (Figs. 14 — 19). The nematocysts of the mesenterial
filaments are about '033 mm. in length, and are found in every stage of development.
They differ from the tentacular nematocysts in that they are relatively much broader,
and are from the youngest stage of about the same size as the ripe nematocvst.
The thread is much thicker with fewer coils, and its lower part, when ejected, is
formed by an eversible portion of the cell. The structure of the ripe uematocyst
(Fig. 14) is very similar to that of the nematocysts from the stomodoeum of Euphi/llia,
described and figured by Bourne (4), and those of Caryoplnjllia by Mobius (20) and
Iwanzoff (17). The eversible portion is never so long as in Curyophyllia, and is not
indeed usually more than about one-third the length of the whole cell (Figs. 14 and 17).
The end of the thread projects in the centre of this to the somewhat pointed extremity
of the cell. Ai'ound the eversible base is a single spiral row of short hairs, with about
OF COENOPSAMMIA FROM LIFU. 371
fourteen turns instead of three rows as described in Caryophyllia. I could not dis-
tinguish in the thread any spiral band as found in Canjophyllia, nor any peculiar
armature at the tip as in Euphyllia. The thi-ead is very thick, and lies in the
ripe condition close under the exterior membrane, which it may bulge out slightly
(Fig. 14). It is coiled from 8 to 10 times on itself, and ends below in a granular
mass of protoplasm at the base of the cell. The latter lies at some small distance
from the structureless membrane in a protoplasmic bed connected by one or more fibres
with the nervous layer.
It is e.xtremely difficult to see the mode of eversion of the nematocyst, but
generally the thread appears to be extruded first, and to carry with its lower end the
eversible base (Fig. 15). Sometimes, the latter can, when the thread is partially
e.Ktruded, be seen to be partially everted, and in one case I found it completely
everted with the thread quite distinctly visible in the middle (Fig. IG). It does not,
however, in any way approach the condition described and figured by Mobius (20),
for the everted nematocysts of Caryophyllia. When the thread is partially, or but
recently extruded, there appears to be a distinct cavity left in the protoplasm (Fig. 16).
In sections too the thread appears to have a distinct sheath, which is quite separate
from the eversible basal portion.
Besides these ripe nematocysts, a large number are found having the eversible
base with central thread well marked, the rest of the nematocyst staining very uni-
formly of a light brown colour in iron haematoxylin and eosin (Fig. 17). In a few
of these cells I found a large nucleus near the base, in one with a well marked
nucleolus. A differentiation of the protoplasm into a slightly darker band, running
spirally round the cell is next seen (Fig. 19), and from this up to the fully developed
thread every stage is clearly visible. The darker band first appears at the outer end —
in respect to the structureless membrane — of the cell, and the thread develops from
this end towards its base.
In the filaments the nematocyst thread is often found completely extruded with
the cell in situ. In other cases the cell is partially or completely extruded from the
filament, but it is impossible to say how far this may not be due to pressure brought
about by the strong contraction of the polyps. I have been unable to find any
appearance of developing nematocysts other than those previously described, — with the
possible exception of the nematocyst represented in fig. 18 — and it seems to me to
be probable that the thread becomes detached somewhere beyond the eversible base,
which is then retracted, a new thread being formed. In some of the extruded ne-
matocysts the thread seems to have been broken off in this way, but there is no
appearance in the eversible base of myophan bands. In one extruded nematocyst there
is a well marked nucleus at the base, and in several ripe nematocysts I have found
a more darkly staining mass within the thread, but with sections only it is almost
impossible to speak with certainty on such points.
Calicoblastic Ectoderm (c. ect. Fig. 20). The polyp next the corallum is every-
where covered by a thin layer of ectoderm, the calicoblast layer. This is not generally
very well marked, and in sections looks like a thin line of tissue, slightly swollen in
places where the small, round, homogeneously staining nuclei are situated. The nuclei
372 ON THE ANATOMY OF A SUPPOSED XEW SPECIES
do not appear to be especially massed in any position, and near the base of the
polj'ps are very distant from one another.
The layer however appears to be complete, and to separate the structureless
membrane everywhere from the coraUum except at the attachments of the mesenteries,
and dividing walls of the coenosarcal canals outside the calices. Three structureless
membranes join one another in these positions (Fig. 20), and at their junction
a number of small bands are given off, which broaden out at their ends, and
are directly attached to the corallum. In section these bands are seen to be
striated, and the calicoblastic ectoderm between them is especially well developed,
almost completely filling up their interspaces. In some polyps, especially at the bases
of the mesenteries, the striations appear to run for some distance into the structureless
membrane and to be due to fibres, but generally it appears quite structureless.
In oblique sections the bands have the appearance, represented by Sclater (25),
in Stephanotrochus, and appear to have no connection with the structureless membrane.
Sclater believed these blocks to be the calicoblasts, but Bourne (4), and subsequently
Fowler (10), pointed out their real nature. In Coenopsammia they are short and have
rather broad bases of attachment to the structureless membrane of the mesenteries, while
at the attachments of the dividing walls of the coenosarcal canals they are generally very
long, much narrower, where they leave the structureless membrane, broadening out very
greatly at their ends. The spaces between them, in the latter position, are crowded with
nuclei, and they often present in thick transverse sections (10 — 12/i) almost exactly the
same appearance as represented by von Heider (13), for the calicoblasts in the same
position in DendrophyUia, a closely allied genus (12. Taf xxxi. Figs. 8, 9 and 11).
I have too very carefully examined the calicoblasts, together with their attaching bands
from the same position in undecalcified and partially decalcified preparations of
Coenopsammia and other Madreporaria, and I can confidently state that neither in them
nor in the calicoblasts are there any crj'stals of any sort. In decalcified preparations of
Pocillopora (5 species), Seriatopora, Prionastraea, Madrepora and C'oenopsaiuinia, I have
found no trace of any organic tissue or remains in the corallum, other than that due
to Clione or boring algae, and there does not seem to be any room for doubting von
Koch's conclusion (18), that the corallum lies completely outside the animal, and is the
result of secretion by the calicoblastic ectoderm, the elements of which retain their cnvn
organic existence.
Endoderm. (end. Figs. 5, 20 — 22). I found it impossible in the hardened and
preserved specimens to tease out separate cells from the endoderm, and in sections no
cell outlines can be seen. In the coenosarcal canals and generally in the contracted polyp
above the tentacles and over the corallum, the endoderm consists of a ragged much
vacuolated epithelium of a cubical facies with large round granular nuclei with distinct
membranes together with a few scattered mucous cells (Fig. 5). Over the muscles and
between the attachments of the mesenteries to the external body wall it has a very
elongated columnar facies with small oval or round, deeply and homogeneously staining
nuclei with a few of the larger granular nuclei (Fig. 21). The structureless membrane
is irregularly drawn out under the endoderm into processes, to which fibres from the
endoderm appear directly to be attached, one or more corresponding to each nucleus.
OF COENOPSAMMIA FROM LIFtJ. 373
No trace of a definite nervous layer can usually be distinguished, except at the
attachment of the mesenteries to the body wall under the tentacles, where it is some-
times well marked with numerous small round nuclei.
The great mesenteric retractor muscles (Fig. 21) are situated on simple or branched
lamelliform folds of the structureless membrane. They consist of separate long fibres,
rectangular or oval in transverse section, and stain very deeply and uniformly with
Heidenhain's iron haemato.xylin. Each fibre runs from the mouth-disc to its insertion
along the line of the attachment of the mesenteries as shown in Figure II. At each
end the fibres bi'eak up into a number of small fibrils, which spread out and appear
to be attached to the structureless membrane without any connection with the endoderm
epithelium. At the attachment of the mesenteries to the corallum some of these
fibrils run directly into the structureless membrane, and I would suggest that the fibres,
previously described in this position, are really the attaching fibrils of these muscles.
The circular rrtuscle — Rotteken's muscle — consists of similar but much smaller fibres
(Fig. 5), the attachment and course of which I have, however, been unable to satis-
factorily determine.
On the sides of the mesenteries opposite to the great retractor muscles, I have
been unable to distinguish any similar muscular fibres running transversely to the long
axis of the polyp — any protractor fibres in fact. In sections numerous small muscular
processes are cut across, but these are quite short, and are basal processes of the
endodermal epithelium. Further, they do not seem to run markedly more transversely
than longitudinally. As the polyp is very closely attached to the corallum, they
could not either have any powerful protractor action, and it seems to me that the
expansion of the polyps must be brought about by other means.
Generative Organs. I have not been able to trace the inwandering of the
genei-ative cells from the endoderm into the structureless membrane, but there is little
doubt that it occurs in the same way as the Hertwigs have described in the Actiniaria
(14 and 15). The endoderm round each ovum is very much thickened, and in the
contracted condition of the polyp the coelenterou between the mesenteries is almost
completely obliterated (Fig. 22). No cell outlines can be distinguished, but the endoderm
appears to be a very elongated and much vacuolated epithelium of a columnar facies.
The nuclei, which are small and not very distinct, are situated towards the free side of
the epithelium, which is very granular. Near the base of the epithelium a few granules
are found, which in places appear to be passing into the ova, large granules bemg
constantly seen situated partially in the ova and partially in the endoderm.
The structureless membrane completely surrounds each ovum as a very thin layer
and the membrane connecting it with the general structureless membrane of the
mesentery is exceedingly fine. It can usually be distinguished, however, by the well
marked basal muscular processes of the endoderm, which are not generally found over
the ova. I have not found any definite " Fadcnapparat " as described by the Hertwigs
in Actiniaria, nor is there any trace of any of the ova serving for food to the rest.
The ova vary up to -45 mm. in diameter and are full of yolk spherules and
granules. The nucleus is a large centrally situated body with a homogeneous deeply
staining round nucleolus. No definite nuclear membrane can be distinguished. Towards
W. IV. ^^
374 ON THE ANATOMY OF A SUPPOSED NEW SPECIES
the exterior under the structureless membrane there is commonly found a thin layer
of granules exactly similar in appearance to the granules found at the bases of the
overlying endoderm.
SECTION IV.
Conclusions relating to the Bodv Layers in the Actinozoa.
The central glandular elements of the mesenterial filaments have been shown by
the Hertwigs (14) and M'Murrich (22) for the Actiniaria, and E. B. Wilson (27) and
Hickson (16) for the Alcyonaria to be the elements, which produce the digestive
secretion. Similar scattered glandular elements have been recorded by most observers
in the stomodoea of Actiniaria and iladreporaria, and recently hav^e been found also
by Ashworth (1) in the stomodoeum of Xenia. The abundance and regular arrangement
of the glandular elements, found in this species of Coenopsammia has however not
previously been observed. In longitudinal sections through the mesenterial filaments
and stomodoeum it is impossible to tell where the latter ends and the former com-
mences so complete is the resemblance between the epithelia of these two parts. The
epithelium too of the lateral parts, the " Flimmerstreifen " is precisely similar to that
of the stomodoeum between the attachments of the mesenteries. It is important also
to remember that the mesenterial nematocysts are found on all the three parts of
the filaments and also in the stomodoeum and external ectoderm of the body.
It was suggested first by E. B. ^yilson (27) that the lateral parts or "Flimmer-
streifen " of the Actiniarian mesenterial filament are ectodermic in origin and that the
median part, or " Drlisenstreifen," is endodermic. Fowler (7) however from the histology
of the Madreporarian filaments considered the median part to be ectodermic and the
lateral parts to be its unbroken gradation into the endoderm but mainly endodermic.
H. V. Wilson (28) then from the development of Manicina came to the conclusion
that the whole filament is ectodermic in origin. Indeed he showed that the filaments
of the twelve primary mesenteries are formed as outgrowths from the basal end of
the stomodoeum. M'^Murrich (22) would not admit the homology of the lateral parts
of the Madreporarian filaments with the "Flimmerstreifen" of the Hert\vigs and main-
tained the views of E. B. Wilson.
In this species of Coenopsammia it is obvious that if the central part of the filament
is endodermic, a great part of the wall of the stomodoeum is likewise endodermic.
Ashworth (1) however states that in Xenia, where digestive cells also occur in the
stomodoeum, he has followed the development in the bud and that the stomodoeum
is entirely ectodermic. I have also found that the stomodoeum is entirely ectodermic
in its origin in the buds of Pocillopora. In this species of Coenopsammia the glandular
elements are found right up to the mouth of the stomodoeum and in the bud
formation, so far as I could follow it, the whole stomodoeum appears to be formed
by the ectoderm. The lateral parts of the mesenterial filaments are similar in structure
to the "Flimmerstreifen" of the Hertwigs and have apparently the same function.
OF COENOPSAMMIA FROM LIFU. 375
They too are directly continuous with the stomodoeum and have the same structure,
so that fi-om the histology one must come to the conclusion that the whole filament of
the pnmary and secondary mesentenes is ectodermic in origin. It would appear also
most probable that the filaments of the tertiary mesenteries are likewise ectodemiic.
I have already pointed out that the central glandular elements of the mesenterial
filaments have been shown to produce the' digestive secretion in both the Actiniaria
and in the Alcyonaria. In one polyp of Coenopsammia, which I have examined by
transverse sections, a small Crustacean is lying in the coelenteron, where it passes
into ■ the stomodoeum. It is noticeable that in spite of the strong contraction of the
polyp it is supported by the mesenterial filaments alone. Willem (29) too has shown
in several Actiuians that the prey is always clasped by the mesenterial filaments after
passing through the stomodoeum, and further has investigated the action of the
digestive secretion op proteids, glycogen and fats. Both Hickson for Alcyoniwm diyi-
tatum and Willem for various Actinians have brought forAvard negative evidence ti>
show that no particles of food are taken up in the solid form by the so-called
endoderm. Particles of carmine are however readily seized and the chief excretive
functions cippear to lie in this epithelium. There are no secretory digestive cells in
the so-called endoderm, and it follows hence that digestion must be brought about
by the ectoderm of the stomodoeum together with its downgrowths over the edges of
the mesenteries, forming their filaments.
The stomodoeum of Zuantharia and necessarily also of Ailcyonaria is not comparable
then to the stomodoeum of the Triploblastica but rather is, with the mesenterial filaments,
the homologue of the ivhole gut. The so-called endoderm, giving rise to the muscular bunds
and generative organs and performing also the excretory functions, is then homologous tuith
the mesoderm of Triploblastica. In the terms of the layer theory, of whatever value it
may be, the Actinozoon polyp must then be regarded as also a Triploblastic form having
definite ectoderm, endoderm and mesoderm.
Sedgwick (26) in 1884- pointed out the possible importance of considering the
Actinozoon polj'ps in connection with the origin of metameric segmentation in Triplo-
blastica, a view which was afterwards strongly supjjorted by van Beneden (2) from
his researches on the development of Arachnactis. The foregoing facts seem to me
to give a strong support to this hypothesis. It is however beyond the scope of this
paper to discuss either this question, or that of the relationship of the Actinozoon
and Hydrozoon polyps.
51—2
376 ox THE AXATOMY OF A SUPPOSED NEW SPECIES
LITERATURE.
1. AsHwoETH, T. H. "The Stomodoeum, Mesenterial Filaments and Endoderm of Xenia.
P. R. Soc. London, Vol. lxiii., p. 44.3, 1888.
2. Bexeden, Ed. van. " Recherches sur le Developpement des Arachnactis." Bull. Ac.
Belgique, 3"= Sen, Tome xxi., p. 179, 1891.
3. Bourne, G. C. " The Anatomy of the Madreporarian Coral Fungia." Quart. J. Micr.
Sci., Vol. XXVII., p. 293, 1887.
4. Bourne, G. C. '■ The Anatomy of Mussa and Euphyllia and the Morphology of the
Madreporarian Skeleton." Quart. J. Micr. Sci., Vol. xxviii., p. 21, 1888.
5. Duncan, P. Martin. " A Revision of the Families and Genera of the Sclerodermic
Zoantharia, Ed. and H., or Madieporaria." J. Linn. Soc, ZooL, Vol. xvill.,
p. 1, 1885.
6. Edwards, Milne and Haijie, J. " Monographic des Eupsammides." Ann. Sci. Nat.,
Ser. 3, T. x., p. 65, 1848. And " Histoire des Coralliaries." 1860.
Fowler, G. H. "The Anatomy of the Madreporaria."
7. I. " Flabellum, Rhodopsammia." Quart. J. Micr. Sci., Vol. xxv., p. 577, 1884, and
Stud. Owens Coll., Vol. I., p. 243, 1886.
8. II. '■ Madrepora." Quart. J. Micr. Sci., Vol. xxvii., p. 1., 1887, and Stud. Owens Coll.,
Vol. II., p. 1, 1890.
9. III. "Turbinaria, Lophohelia, Seriatopora, Pocillopora." Quart. J. Micr. Sci., Vol. xxviii.,
p. 1, 1888, and Stud. Owens Coll., Vol. ii., p. 17, 1890.
10. IV. "Madracis, Amphihelia. Stephanophyllia." Quart. J. Micr. Sci., Vol. xxviii., p. 413,
1888.
11. V. " Duncania, Madrepora, Galaxea, Heteropsammia, Bathyactis." Quart. J. Micr. Sci.,
Vol. XXX., p. 405, 1890.
12. Hamann, O. " Der Organismus der Hydroidpolypen." Zeitschr. Naturw., Bd. xv.,
p. 473, 1881.
13. Heider, a. R. von. " Korallenstudien." Zeitschr. wiss. Zool., Bd. xliv., p. 507, 1886.
14. Hertwig, O. und R. "Die Actinien." Jena, 1879.
15. Hertwig, R. "The Actiniaria." Challenger Reports, 1882. Suppl., 1888.
OF COENOPSAMMIA FROM LIFU. 377
IG. HiCKsox, Sydney, J. "The Anatomy of Alc3'onium digitatura." Quart. J. Micr. Sci.,
Vol. XXXVII., p. 343, 1895.
17. IwANZOFF, N. "Ueber den Bau, die Wirkungsweise und die Entwickelung der Nessel-
kapseln der Coelenteraten." Bull. Soc. Moscou, 1896, p. 1.
18. Koch, G. von. "Ueber die Entwicklung des Kalkskeletes von Astroides calycularis und
dessen morphologische Bedeutung." Mt. Stat. Neapel, Bd. iii., p. 284, 1882.
19. Lendexfeld, R. von. "The Functions of Nettle Cells." Quart. J. Micr. Sci., Vol. xxvii.,
p. 393, 1887.
20. MoBius, Karl. " Ueber den Bau, den Mechanismus und die Entwicklung der Nessel-
kapseln einiger Polypen und Quallen." Abh. Ver. Hamburg, Bd. v., p. 1, 18GC.
21. MosELEY, A. N. "Report on the Corals." Challenger Reports, 1881.
22. McMuRRiCH, J. Playfair. " On the Development of the Hexactiniae." J. Morphol.,
Vol. IV., p. 303, 1891.
23. Ogilvie, ilaria M. " Microscopic and Systematic Study of Madreporarian Types of
Corals." Phil. Trans., Vol. clxxxvii., p. S3, 1896.
24. Ort.mann, a. " Beobachtungen an Steinkorallen von der Siidkuste Ceylons." Zool.
Jahrb., Bd. iv., SuppL, p. 493, 1889.
25. ScLATER, W. L. " On a Madreporarian Coral of the Genus Stephanotrochus from the
British Seas." P. Zool. Soc. London, p. 128, 1886.
26. Sedgwick, Adam. " On the Origin of Metameric Segmentation and some other Morpho-
logical Questions." Quart. J. Micr. Sci., Vol. xxiv., p. 43, 1883.
27. Wilson, Edmund B. "The Mesenterial Filaments of the Alcyonaria." Mt. Stat. Neapel,
Bd. v., p. 1, 1884.
28. Wilson, Henry V. " On the Development of Manicina areolata." J. Morphol., Vol. ii.,
p. 191, 1889.
29. WiLLE.M, Victor. " La Digestion chez les Actinies." Bull. Soc. Medecine de Gand,
p. 295, 1892.
378 ON THE ANATOMY OF A SUPPOSED NEW SPECIES
EXPLANATION OF PLATE XXXIV.
Lettering used througJiout.
ect. Ectoderm. end. Endoderm. s. tn. Structureless membrane or lamella. c. ect.
Calicoblast ectoderm. n. I. Nervous layer of the ectoderm. m. g. c. Mucous gland cells.
g. g. c. Granular gland cells. 7n. n. Mesenterial nematocysts. t. n. Tentacular nematocysts.
mus. Muscular fibres, ov. Ovum. m. J. Mesenterial filaments. »i. 1 Primary mesentery.
m. 2 Secondary mesentery, m. 3 Tertiary mesentery.
Fig. 1. View of a large colony from the side (cleaned corallum). Nat. size.
Fig. 2. A small colony from above. One parent and three daughter corallites. Nat. size.
Fig. 3. A single calice from above. 1, 2, 3 Primary, secondary and tertiary septa.
Fig. 4. Diagrammatical section of the ectoderm of the polyp outside the tentacles.
Mucous gland cells {m. g. c.) are everywhere numerous, but the granular gland cells
(g. g. c.) are only found near the base of the tentacles. Two kinds of nematocysts are
found, of which the mesenterial form («i. n.) is much the less numerous. The granular
nervous layer {n. I.) is not well marked except at the bases of the tentacles ; in it a few
large nuclei of nerve cells can commonly be distinguished.
Fig. 5. Section through the body wall at the sphincter muscle, the fibres (mus.) of which
are small and flattened. No cell outlines can be distinguished in either the ectodermal or
endodermal epithelia.
Fig. 6. Section through the middle of a single battery of nematocysts on one of the
primary tentacles. The central part of the battery is packed ^-ith nematocysts under which
the nervous layer is very conspicuous, while at the sides the ectoderm cells end in muscular
processes on the structureless membrane.
Figs. 7 — 10. Tentacular nematocysts. (Oc. 4, oil immersion ~.)
Fig. 7. A ripe tentacular nematocyst with the thread fully formed. Outside the basal
end is a conspicuous oval nucleus in a finely granular mass of protoplasm, which forms the
granular (nervous ]) peduncle.
Fig. 8. A ripe tentacular nematocyst with the thread partially extruded, found lying
freely in the cavity of one of the retracted tentacles.
Fig. 9. Developing nematocyst. In place of the extruded nematocyst a homogeneous
mass of protoplasm appears, the central part of which acquires a definite membrane and
becomes finely granular.
OF COENOPSAMMIA FROM LIFU. 379
Fig. 10. Later stage tlian Fig. 'J. The young nematocyst decreases in size and becomes
coarsely granular, the granules arranging themselves in a spiral line close under the external
membrane. The nematocyst drawn has been cut rather obliquely so that in the upper half
the granules of the lower side are seen, and in the lower half those of the upper side, the
granules in the centre appearing to be almost irregularly arranged. A comparison with
Fig. 7 will make this clear.
Fig. U. Transverse section through the stouiodoeum sliowing the attachment of three
mesenteries. The epithelium of the stomodoeum is very thick — being formed apparently of
cells of an elongate columnar faoies — and opposite to the attachments of tlie mesenteries
crowded with goblet-like vacuoles.
Fig. 12. Transverse section through a primary mesenterial filament immediately before
its convolutions commence. The structureless membrane ends in a T-shaped expansion in
the filament, which is well marked ofi" from the general endoderm of the mesentery. It is
distinctly divided into three parts, a central, crowded with goblet vacuoles, and two lateral,
crowded with homogeneously staining oval nuclei. A single nematocyst is seen in the central
part, but the nematocysts do not become numerous until its lower half is reached.
Fig. 13. Transverse section of the same filament as in Fig. 12 in its lower third. The
filament is as distinctly marked off from the endoderm of the mesentery below it, but it is
no longer divisible into three parts, being crowded with nematocysts in different stages of
development — one with the thread extruded and cut off short.
Figs. 14 — 19. Mesenterial nematocysts. (Oc. 4, oil immersion xV ** f-)
Fig. 14. A ripe nematocyst. The thread ends below in a mass of granules at the base
of the cell while the opposite end appears to lie freely in an eversible sheath, marked by
fine lines due to a spiral row of fine hairs. (Somewhat diagrammatical.)
Fig. 15. Nematocyst with part of the completely ejected thread. The thread seems to
be extruded first and to carry behind it its eversible base, which is surrounded by a distinct
row of spiral hairs. At the upper end of the nematocyst, round the eversible base of
the thread, is a distinct depression.
Fig. 16. Nematocyst with ejected thread, whicii is still however visible in the middle of
the basal portion. In the body of the nematocyst traces of a spiral sheath can be seen, from
which the thread seems to have been ejected.
Fig. 17. Nematocyst with eversible base but without any appearance of a thread in
the cell. Near the base is a nucleus with nucleolus (both seen very rarely) and the cell
terminates in a nucleated peduncle, which bi-anches out over the structureless membrane.
Fig. 18. Nematocyst possibly in an earlier stage to Fig. 17, without any distinct base
for the thread and with a distinct nucleus and nucleolus.
Fig. 19. Nematocyst of a later stage to Fig. 17, with a differentiation of the protoplasm
into a dark spiral band, which will subsequently become the thread, lying in a clearer areji.
Fig. 20. Section through the base of one of the dividing walls of one of tlie coenosarcal
canals showing the attachment of the structureless membrane to the corallum. The foi-nier
is drawn out into long striated bands, swelling out at their ends where they are attached
to the corallum. The striations appear in some cases to be due to fibres in the structureless
membrane, but they are not generally so well marked as in this section. The spaces between
the bands are completely filled by the calicoblast ectoderm.
380 ON THE ANATOJIY OF A SUPPOSED NEW SPECIES OF COENOPSAMMIA.
Fig. 21. Transverse section through a portion of a longitudinal retractor muscle of a
primary mesentery. The structureless membrane is drawn out into simple or branched
lamelliform folds, on which the somewhat rectangular or rounded fibres are placed.
Fig. 22. Transverse section through a primary and a secondary mesentery at a slightly
lower level to that represented in the left half of sextant F in Figure I. The coelenteron is
almost obliterated. The tertiary mesenteries {in. 3) are recognisable as fine bands. Neithei-
the primary nor secondary mesenteries have any trace of muscular bands but in the
structureless membrane of each an ovum is situated. Round these the endoderm is much
thickened ; its nuclei are generally indistinct but towards the free surface it is crowded with
fine granules. Over the structureless membrane a few larger granules are found and can be
seen in every stage of their passage into the ova, where they form a row of granules round
the periphery. The nucleus of the ovum is large with large nucleolus : no nuclear membrane
however is generally \'isible.
Zoological Laeoratort,
January 31, 1899.
■■>",»
\-^r-it-r-t
^^'^'^
%
.-at]
J.SG.ani Edf/in Wilson dei
nARniNF.R. COENOPSAMMIA,
ON THE INSECTS FKOM NEW BRITAIN.
By D. sharp, M.A., MB., F.R.S.
With Plate XXXV.
The insects obtained by Dr Willey in Xew Britain and Lifu have ah-eady been
the subject of a paper in the first part of these " Results'." The specimens of
hexapodous Insects brought back by Dr Willey are fairly numerous. Except in Coleo-
ptera and Orthoptera they are all from New Britain. The number of species in com-
parison with the number of the specimens is very large, so that the collection is not
of a kind that it is desirable to work out in complete detail. They show that New
Britain is rich in insects, for unless this were the case it is impossible that Dr Willey
should have obtained so many species as he did ; entomological research having been
only subordinate to some other objects of his expedition ; besides which it must be
recollected that his researches were confined to one or two localities on the coast.
Very little work has at present been published on the Entomology of New Britain.
And even in the case of New Guinea — the larger island of which New Britain is so
near a neighbour — the fauna has only been very imperfectly ascertained. In these notes
I propose only to touch on certain species that can be advantageously dealt with.
As no allusion is subsequently made in this paper to certain of the Orders of
Hexapoda I may here mention that the Neuroptera are represented by very few
specimens, and that the Hemiptera have been entrusted to Jlr G. W. Kirkaldy for
examination. The latter are extensive in comparison with the other Orders, and it is not
probable that Mr Kirkaldy's account of them will be ready in time to be included
in these " Results." Lepidoptera are represented in the collection by only a few larvae
and pupae^.
' " Account of the Phasmidae, with notes on the Eggs," pp. 75 — 94, Plates VIII. and IX. Dr M. v. Brunn
has since published (Mt. Mus. Hamburg, xv., 1898, p. 4) a note on the egg referred to on p. 89 and PI. IX.
f. 39 as that of a Ci/phocraiiia without name, and for which the name of C. lianiUchi was suggested.
According to Dr v. liruun, tliis insect is really the Cyphucraiiiii herculeana of Charpentier, Westwood having
been wrong in treatin;; Charpentier's species as a synonjm of C. goliath. Herr Brunner v. Wattenw^l, in
his expected monograph of Phasmidae, will separate these two species as a genus with the name Euri/ciiema
Serv. : the name of this Phasmid will therefore be Eunjcnemu herculeana Cliarp.
- An account of the butterflies of New Britain has recently been published by Herr C. Bibbe in Deultche
ent. ZeiUchr. Lep., 189M, pp. 3.0—133.
w. IV. 52
382 ox THE IXSECTS FROM NEW BRITAIN.
Order Coleoptera.
The collection of Coleoptera obtained by Dr Willey in New Britain numbers about
160 species. In 1883 Fairmaire published an essay on the Coleoptera of the New
Britain archipelago' iu which he enumerated 176 species. Since then but little has
been added. Upwards of one half of the species mentioned by Fairmaire are not
represented in Dr Willey 's collection ; moreover the majority of the species enumerated
by Fairmaire really came from Duke of York island.
In the following remarks I have mentioned onlv such names as add to the
knowledge supplied by Fairmaire's paper. Many of the species procured by Dr Willey
are clearly undescribed, but I do not think it desirable to deal with the obscure
forms that are represented only by one or two specimens, or with others that cannot
for various reasons at present be satisfactorily elucidated.
Dr Willey also procured a small collection of Coleoptera in Lifii. This is at
present being studied by M. Albert Fauvel of Caen, who is well acquainted ■ivith the
Coleoptera of New Caledonia.
Fajiilt LUCANIDAE.
Eurytrachelus intermedius Gestro, Ann. Mus. Genova, xvi., 1881, p. 317.
Plate XXXV. Figs. 3 a, 3 6, 3 c, 3 rf.
Dr Willey procured a good series of this rare stag-beetle in the neighbourhood
of Blanche Bay ; nine specimens are of the teleodont form (Plate XXXV. Fig. 3 a), two
of the mesodont form (Fig. 3 6), and two of the priodont (Fig. 3 c). There are also
four females (Fig. 3 d). The forms of the male, besides differing in the mandibles,
exhibit great variation in the sculpture. Gestro described the species from New Guinea,
and Dr Willey's examples differ a little from Gestro's figure ; chiefly in that the
mandibles are more slender and their teeth less largely developed. A specimen in
the British Museum, reputed to be from Duke of York island, quite agrees with the
New Britain specimens. E. intermedius is closely allied to E. ternatensis. The latter
species is recorded by Fairmaii-e from Duke of York island, but we may anticipate
that this record will prove erroneous : E. intermedius having been probably mistaken
for E. ternatensis.
Family SCARABAEIDAE.
Phaeochrous alternatus Fairm. (t. c. p. 5). Several specimens of this beetle were
taken from the stomach of Varanus indicus. It was not otherwise noticed.
? Xylotrupes gideon L. Plate XXXV. Figs. 1, 2.
Xylotrupes gideon is recorded from Duke of York island by Fairmaire, without
any remark as to its variation. The four specimens of a male Xylotrupes procured
by Dr Willey represent various grades of development of the horns of that sex, and
' Ann. Soc. ent, Belgique, xxvn., 1883, Pt. II., pp. 1 — 58.
ON THE INSECTS FROM NEW BRITAIN. 383
are of some interest as they exhibit no sign of the secondary projection on the
lower horn, that is invariably found on the specimens of A'', gideon from India and
Java. It i.s not possible in the present state of our knowledge to form an opinion
as to whether these examples are a different species or not. M. van Lansberge has
described a Dynastid fi'om the island of Flores (Endebiiis florensis) the male of which
is said to be very similar to that sex of Xylotrupes gideon, while the female is so
different that van Lansberge has proposed a distinct genus for the species. Dr Willey
brought back only one female ,that could possibly be the other sex of the male we
are discussing, but there is no certainty that it is so, and if the two insects are
really the sexes of one species, it is clearly not X. gideon.
We have figured the profile of the head and thorax of a Javanese example of
X. gideon (Plate XXXV. Fig. 2) for comparison with our New Britain insect.
Family MALACODERMIDAE.
There have long been known to entomologists some extremely remarkable larvae
that probably are those of Lampyrides, or Lycides, though none of them have been
satisfactorily identified '.
Dr Willey procured a most remarkable form of this kind, bearing long abdominal
processes, that are segmented or articulated at the base (Plate XXXV. Figs. 4, 4 a,
4 b). I take this opportunity of drawing attention to these forms with the hope that
some one may soon be able to give us some further information about them. Although
the larvae I have mentioned as being presumably those of Lamp}Tides or Lycides, are of
very diverse forms, yet they all have a head with well-developed mouth-parts, and capable
of being retracted into the tubular cavity placed underneath the shield-like prothorax.
They have also well-developed prothoracic legs terminated by a single claw. It is probable
that they may prey on Mollusca.
Family CERAMBYCIDAE.
Arrhenotus willeyi, n. sp. Plate XXXV. Fig. 5.
Niger, parum nitidus, pube albicante variegatus, in elytris fasciis duabus flammulatis;
prothorace, utrinque biaugulato, tuberculo inframarginali, elongate, acuto. Long. iO —
2-5 mm.
Head a good deal narrower behind the eyes, sparingly and in-egularly punctate,
variegate with a few white spots. Thorax strongly transverse, a good deal narrower
than the elytra, dorsum a little uneven, irregularly and variably spotted w itii white :
on each side the margin forms in front a well-marked ])rominent angle, and in the
middle a much more obscure obtuse angle ; below the margin there is an acute spine,
white above, black below. Scutellum white on each side, black in the middle. Elytra
much pnjduced on each side of the scutellum, and with a small angle projecting inwardly
' It appears probable tliat n fossil larva of this group has buen mistaken for a Crustacean allied to
Apodidae. Cf. (iahan, Xaturnl Seifm-i', xii., 1898, p. 42.
52—2
384 ox THE INSECTS FROM NEW BRITAIN.
from the front part of the ba}' formed by their prominence ; base with a short carina
projecting angularly at the shoulder : their surface marked with iiTegular and variable
white spots, the largest of which are combined to form two transverse bands, very much
indented, the anterior band is directly transverse, the posterior one strongly angulate,
projecting forwai'ds at the suture ; behind these two bands some irregular white spots :
the sculpture variable, there being some subseriate punctures, and the interstices being
more or less longitudinally raised : the punctures at the basal parts of the sides are very
numerous and coarse : tips truncate, each \vith a broad short spine externally. Under-
surface, irregularly variegate with white.
The three individuals of this species are probably two of them female, one male ; if
so there is very little difference between the sexes.
Thomson and Pascoe established several genera for the allies of Arrhenotus which
were subsequently treated by Lacordaire as mere sections. A. luilleyi does not enter
satisfactorily into any of these sections. In coloration and general appearance it some-
what resembles Elais exarata Pascoe. (Trans, ent. Soc. London (3) III., PI. XIX.
Fig. 7.)
Tmesisternus yorkensis.
SpJnngnotus yorkensis Fairmaire, Xaturaliste, 1881, p. 359: Ann. Soc. ent. Belgique,
XXVII., 1883, p. 47.
I refer a series of examples to this species notwithstanding the fact that they
belong to the genus Tmesisternus, not Sphingnotus, and that they present some differ-
ences from Fairmaii-e's description. The punctures of the el}-tra are not serially disposed ;
the apical markings of the elj^ra and those on the abdominal segments are variable.
The series procured by Dr Willey consists of 20 males and 12 females. The size varies
fr-om a length of 19 to one of 27 mm., and this variation occurs equally in the two
sexes. The peculiar swelling of the middle tibiae of the male also varies greatly, as
does the sculpture on the anterior aspect of the swollen part. The males and females
are very much alike, but can be invariably distinguished by the last ventral plate.
This is longer in the female and is more or less longitudinally impressed along the
middle, the impression behind becomes much broader. The male has no trace of this
impression, and has more abundant dark hair on the apical part of the segment. In
both sexes the hind angle of this plate projects back as a short spine, longer in the
female than in the male.
Fairmaire's specimens were from Duke of York island. Not suspecting at first
that Dr Willey's specimens belong to Fairmaire's species, I had proposed to give them
the name of Tmesisternus tardus. Dr Jordan has described an allied form from New
Guinea under the name of T. dohertyi, Nov. Zool. I. p. 500, and remarks that it is
near T. yorkensis Fairm. ; it was this remark that led me to the above identification,
which I believe will prove to be correct.
Diochares basigranatus Fairin. (t.c. p. 51). I think Fairmaire is correct in con-
.sidering this distinct from D. fimbriatus. Dr Willey obtained two specimens of D.
basigranutus. Fairmaire records (I.e.) D. fimbriatus from Duke of York island.
ON THE INSECTS FROM NEW BRITAIN. 385
Monohammus fasciatus Montrouzier (Ann. Soc. Agric. Lyon, vii., 18.5-5, p. G3).
Two specimens, agreeing with others from Woodlark island and New Guinea in my
collection. Fairmaire records three other species of the genus from Duke of York island.
Dr Willey obtained seven specimens, belonging possibly to four different species, near
Blanche Bay, but this material is not adequate for the description of new species in
the very difficult group of M. xylotrephes, to which these forms belong.
Batocera lactiflua Fairm. (t.c. p. 50). Fairmaire's type was a female, entirely white
in colour. Dr Willey has obtained a single female of this peculiar species, but it is
of a griseous colour, deeply suffused with pink.
Batocera nehulosa Bates (P. Zool. Soc. London, 1877, p. 158). Recorded both by Bates
and Fairmaii'e from Duke of York island : now obtained by Dr Willey in New Britain.
Xiphotheata luctifera Fairm. (Le Naturaliste, 1881, p. 359) PL XXXV. Figs. 6, 6 a,
6 6; Ann. Soc. ent. Belgique, xxvii., 1883, p. 49. There appear to be only two species
known of this rare and peculiar genus. Fairmaire grounded his species mainly on the
fact of the male being destitute of the peculiar long horn on the front coxae, a
character that is very conspicuous in X. saundersi. Dr Willey procured three males
and one female of a Xiphotheata that I refer to Fairmaire's species. They show that
the male character mentioned above is merely an individual one; the horn being present
and very largely developed in two of these males (Plate XXXV. Fig. 6), and only a
short spine in the other (Fig. 6 a). The female is easily distinguished by the entire
absence of armature on the front tibia (as well as by the unarmed coxa, Plate XXXV.
Fig. 6 b) and by the terminal ventral plate not being prolonged at the sides behind.
Faii-maire's species may however be maintained, as the colour and sculpture are a little
different from those of X. saiuidersi. This latter species was found by Wallace in
Batchian, Morty, and Gilolo. Fairmaire's specimen was from Duke of York island.
Serixia longicornis Pascoe (Tr. ent. Soc. London, 3rd ser. ill. p. 339). Two specimens.
Previously recorded from Singapore, Ceram, Batchian, Bourn, and (with doubt) Waigiou.
Glenea extreina, n. sp. Rufotestaceus, antennis nigris ; capite thoraceque ochraceo-
tomentosis, hoc medio macula nigra ornato; elytris cyaneis, tomento griseo obscuratis,
apice truncato, singulo angulo externo breviter spino,so, interno recto; pectore abdominisque
lateribus ochraceo-tomentosis. Long. 13 mm.
Antennae slender, rather longer than the body (in the male ?), quite black. Thorax
not quite so long as broad, a little constricted in front of the base. Elytra with
numerous large punctures irregularly arranged and wanting behind, cyaneous, but both
colour and sculpture rendered obscure by a dense, pallid, griseous tomentum ; there are
numerous erect hairs at the shoulders : the humeral angles are sharply marked, fi'om
each there extends backwards a carina that becomes obsolete before reaching the apex :
very near to this more dorsal carina there is a second one that does not commence
at the base, but becomes more definite behind, and projects .so as to form the terminal
spine : the epipleural margin is also strongly raised : the sculpture on the pseud-epipleurae
is very coarse, and the purple (or violet) colour is not obscured by tomentum as it is on
386 ox THE INSECTS FROM XEW BEITAIX.
the dorsum. The claws are very strongly toothed at the base. Four specimens, very
similar to one another, and probably all males. The species appears to be nearer G.
aluensis than any other.
Glenea venus Thomson. One specimen.
Family BRENTHIDAE.
Cacoschizus, n. g. (Ceocephalides).
Tarsi quinque-articulati ; articulis 1° et 2° transversis, 3° quadrato, supeme latissime
fere ad basin impresso, margine apicali sat profunde emarginato, 4° 5°que crassis, 4°
paulo ultra apicem tertii extenso, ab quinto bene discreto, hoc ceteris conjunctis
longitudine fere aequali.
I establish this genus for a Brenthid with very peculiar feet. The species I
believe is Schizotrachelus schmeltzii Fairm. Lacordaire in describing the genus Schizo-
trachelus (Genera Col. vii. p. 455) describes the tarsi as follows: " tarses spongieux
en dessous, a articles 1 — 3 courts, egaux, 3 entier." In the structure of the feet
Cacoschizus comes nearest to certain species of the genus Trachelizus, but from that
genus it is readily distinguished b}' the head being separated from the neck by a verv
deep constriction.
Cacoschizus schmeltzii. PI. XXXV. Fig. 7, c/ ; 7 a, $ : 7 b, hind foot of J'.
Schizotrachelus schmeltzii Faii-m., Ann. Soc. eut. Belgique, xxvil., 1883, Part il. p. 44.
Fairmaii-e described the male onlj', we figure the two sexes; in the structure of the
feet the female agrees with the male. The peculiar tarsi are of considerable interest, as
distinctly 5-jointed tarsi are of very rare occurrence in the Rhynchophorous series of
Coleoptera. The resemblance of both the sexes of Cacoschizus to Trachelizus is very
great, and though Schizotrachelus and Trachelizus are widely separated in Lacordaire's
classification, I think they are nevertheless naturally allied.
Cacotrachelus, n. g. (Eutrachelides).
Mas. Caput elongatum, convexum ; rostrum breve, apice latiore, mandibulis brevibus
spatio parvo includentibus. Antennae breves, crassae. Prothorax convexus, subovalis,
anterius utrinque obsolete convexo, dorso posterius subtiliter canaliculato. Pedes breves
crassiusculae, femoribus brevibus, basi hand pedicellato ; tibiis brevibus compressis, apice
interne mucronato ; tarsis crassis, subtus spongiosis, articulato tertio breviter bdobato.
Lacordaire's gi-oup Eutrachelides consists of a single species of gigautic Brenthidae —
Eutrachelus temmirickii — from Java. The genus I am at present establishing is totally
different from Eutrachelus in appearance, but is I think really allied to it. I have long
had in mj- collection a Brenthid from Java that is closely allied to the insect brought
ON THE INSECTS FROM NEW BRITAIN. 387
by Dr Wille}' from New Guinea, aud I will take this opportunity of briefly diagnosing
it'. The coloration of these insects is very unusual, exhibiting as it does the yellow
lines of the South-American Brenthides in a somewhat different form. From a taxo-
nomical point of view Cacotrachelus is of considerable interest, as it might almost as well
be placed amongst the S. American Brenthides as near Eutrachebis.
Cacotrachelus sculptipennis, n. sp. Plate XXXV. Fig. 8 j/^ ; 8 a $ , head and thorax
from side.
Nigricans, capite thoraceque metallescentibus, femoribus tibiisque flavo-rufis ; elytris
rufis, versus suturam piceis, interstitio tertio fere toto, 5° ad basin flavescentibus, crenato-
sulcatis; capite thoraceque canaliculatis, illo ad verticem profunde impresso. Long, cum
rostro, 8 mm.
Male. Rostrum broad and short, not so long as the thorax, thicker at the tip, deeply
sulcate along the middle ; head elongate convex, canaliculate, the channel expanding
behind into a broad deep depression; separated from the neck by a very deep de-
pression ; the back of the head is somewhat depressed, and on each side with a small
notch or fovea. Antennae inserted in the middle of the rostrum, thick and short,
thicker towards the tip; joints 3 — 10 transverse, 9 and 10 distinctly longer than those
preceding them, 11th joint acuminate, rather longer than broad. Prothorax longer than
broad, rounded at the side and naiTowed in front, convex, impunctate, very distinctly
channelled behind, the channel finer in front, and not reaching the anterior margin.
Elytra but little prolonged behind, truncate at the tip, the outer angle slightly obtuse
and the lateral margin strongly raised behind ; externally deeply grooved, the gi-ooves
very regularly sculptured, the interstices narrow, the first and second striae are fine,
and the first, second and third interstices comparatively broad, the sixth interstice
is somewhat more prolonged and raised at the extreme base. The legs are short,
reddish-yellow, the trochanters and tarsi blackish, the knees a little darker; all the
legs provided with strong angular mucro at the tip of the tibia ; tibiae broad
compressed, but less so in the middle than at the base and apex.
Female. Resembling the male, but with the rostrum slender, and only slightly
thicker at the tip, the antennae inserted near the base, and the tibial mucros obsolete.
Ithystenus dehilis, n. sp. Plate XXXV. Fig. 9 ^.
(/. Niger, opacus, femoribus parte basaii rufa, elytris flavo-lineatis, ad apicem acumi-
uibus duobus, brevibus, simplicibus ornatis. Long, cum rostro, 23 mm.
Allied to the New Guineau /. linearis, but smaller, with the yellow lines of the
elytra extending nearly to the tip, and the apical processes, simple points, without lobe
or swelling at their bases. The hind tibiae are remarkably short, but the femora extend
1 Cacotrachelus javamm n. sp. Picescens, pedibus ruBs, piceo-variegatis ; elytris regulariter crenato-sulcatis,
dorso deplauato, apice subprolongato, truncate; externe, suturaque nigricantibius, interstitio tertio flavo, latiore,
interstitiis 2" 4"iiue vage rufescentibus, 5" iterum llavoscontiore ; antennis crassiusculis, modice elongatis,
apicem versus latioribus, articulis ultimis tribus paulo latioribus; tibiis interinediis et poatcrionbus parte
supra medium crassiore. Long. <?, cum rostro, 13 mm. Hab. Willis mounttiins, .Java.
388 ON THE INSECTS FROM XEAV BRITAIN.
slightly beyond the tips of the ehi:ra. Only one specimen was found. The occurrence
of this remarkable genus in New Britain or the Duke of York island has not been
previously noticed.
Order Hymenoptera.
New Britain is evidently rich in Hymenoptera, as the small number of examples
obtained by Dr Willey consists of comparatively many species. They belong chiefly to the
Aculeata. I have not ventured to describe any of them except a species of Thynnus
that possesses a peculiar abdominal structure. I may however allude to an abeiTation of
instinct observed by Dr Willey in the case of a wasp of the genus Polistes (probabh- an
undescribed species allied to P. colonicus). Instead of one egg being placed in each cell,
there are several, as shown in Fig. 14, PI. XXXV., which represents five cells of the
nest of the insect in question. The cell on the left, below, is closed for the purpose of
pupation, and the one above it is empty. The three cells to the right contain, respec-
tively, one .5, one 3, eggs, and the other (the one to the right) 2 just-hatched larvae.
Notwithstanding the supernumerary eggs, only one larva in each cell attains maturity,
though how the others are disposed of we do not know, Dr Willey ha^"ing made no
observations on this point. Neither did he ascertain whether this aberration is com-
mon in the species, or confined to this nest. He brought back the nest of a second,
and smaller, species of Polistes (also probably undescribed), and in this case there is
only one egg in each cell, as one would e.xpect. It does not seem possible to account
for so striking an aberration of instinct as this, by supposing that there were more
eggs produced than cells to place them in, because several of the cells in the nest
are quite empty.
Family APIDAE.
KOPTORTHOSOMA sp.
Koptorthosoma sp. atF. K. aestuantis, Perkins, Ent. Mo. Mag., Feb. 1899, p. 38.
Mr R. C. L. Perkins has (1. c.) called attention to the very extraordinary sym-
biosis of female bees of the genus Koptorthosoma and certain Acai'ids ; the bee being
provided with a special chamber in the abdomen which is tenanted by the Acari.
The males do not possess this structure ; Mr Perkins mentions the remarkable fact
that in this species from New Britain the female is destitute of the special chamber,
though it exists in the closely allied K. aestuans. Dr Willey only procured two
females of this interesting species, and no male.
Fasiilt THYNNIDAE.
Thynnus serriger, n. sp. PI. XXXV., Fig. 13, $ ; 13 a, extremity of abdomen.
$. Nigricans, hie inde pallide setosus; scutello, abdominis maculis lateralibus,
fasciaque mediana in medio interrupta, albicantibus. Long. 12 mm.
ON THE INSECTS FROM NEW BRITAIN. 389
Vertex of head shining, front densely and coarsely punctured and pubescent.
Thorax transversely quadrate, rugose above, with a carina along the middle ; the
small scutellum almost white, very hairy. Propodeum with very little sculpture, abruptly
declivous behind ; the portion in front of the declivity very short. Abdomen with
the tirst segment very deeply impressed in front, at the edge of the impression \ery
hairy, especially in the middle; just behind the large impression there is a small
depressed area, faintly metallic in colour, and coarsely punctured, but both the peculiar
colour and sculpture are much concealed by the dense pubescence of this part. The
second segment is covered above by coarse, transverse wrinkles, the following segments
being smooth and polished. There is a pallid spot on each side of segments 1 — 4 ;
that on the first segment stretches inwards towards its fellow, as does also that on
the third segment; the spots on the 2nd and 4th segments are quite small. The
peculiar plate at the end of the abdomen is terminated by a pair of saws (PI. XXXV.
fig. 13 a).
Only one specimen was obtained of this species. Though in form, colour and
sculpture similar to various other species of the genus, — T. atratus, e.g. — it is remark-
able on account of the pair of saws at the extremity of the abdomen. Thynnidae
are believed to depredate on pupae or larvae of Lepidoptera underground, and it is
possible that these saws may be useful in penetrating cocoons. So little is known
as to the life-histories of these peculiar insects that this suggestion can be considered
as little more than a random guess. Dr Willey did not obtain any male Thynnus.
Order Diptera.
Of this neglected Order of Insects the specimens brought back from New Britain
are few. They were all placed in spirit, and consequently are mostly, since drying,
in a shrivelled condition : this method of collection being unsuitable for insects of
this Order. All the collection was made in New Britain : Dr Willey did not bring
back any Diptera from Lifu.
In addition to the four species I have ventured to describe, the collection in-
cludes about 24 species. There are several Tipulidae of a commonplace character,
looking like European insects. The family Stratiomyidae is represented by a pair of
Engunia consobrinu, and two species of Ptecticus. The Engonia is of considerable
interest on account of the great difference in the structure of the antennae of the
two sexes. The male — which is twice the size of the female — has the intermediate
joints of the antennae distinctly segmented, and the terminal three joints elongate,
and densely hairy ; the terminal joint being remarkably long. In the female the
intermediate joints are closely compacted and swollen, and the terminal joint is short.
The family Therevidae is represented by a new species of Leptipalpus allied to
L. waigiensis Bigot. Asilidae are apparently common in New Britain, a species of
Laphria (Maira) very near to L. cenea Macq. being represented by twelve specimens,
and there are also two or three species of Oinmatias.
w. IV. • 53
390 ox THE INSECTS FROM NEW BRITAIN.
There are two species of Syrphidae in addition to the Einnerus described below.
The Eumyiid lluscids are apparently numerous : the most remarkable being an
apparently new RutiHa of very brilliant colours, but with the abdomen dark. Of
Acalyptratae there are two forms allied to Calobata, and a species of Senopteinna.
Speaking roughly these Insects seem to be allied to both Australian and Malayan
forms.
Family SYRPHIDAE.
Microdon pictipenne, n. sp.
Gracile, nigrum, coerulescenti-micans, hie inde aurato-pubescens, geniculis tarsisque
testaceis; alls elongatis, hyalinis, nigro-pictis. Long. 10 m.m.
Head shining, violet ; the face on each side with a broad line of golden pubescence ;
antennae black, with the basal joint yellow beneath, the third joint reaching nearly
to the mouth, the seta inserted laterally near its base. Thorax violet, dull, bearing black
pubescence, at the sides in front of the wings with golden pubescence ; there are also
some golden hairs on the scutellum and along the sides of the dorsum. Abdomen
slender, shaped like that of a wasp, violet, the sides and hind margins of the segments
with some golden hair. Femora violet, the base of the tibiae and the tarsi yellow,
the front tarsi more obscure yellow. Wings elongate, reaching about to the tip of
the body, transparent, the nervures very strongly marked, black, the apical portion of
the wing with some irregular black marks extending across the vdng. Halteres white.
One specimen.
This species, like others of its congeners, has a pronounced general resemblance
to Hymenoptera.
Eumerus speculifer, u. sp. PL XXXV. Fig. 10, hind-leg.
Niger, subaeneo-micans, abdomine lunulis albidis ornato, antennis geniculis, tarsisque
sordide testaceis. Long. 7 m.m.
if Head black, between the eyes, above the antennae rendered snow-white by a
very fine depressed pubescence : eyes meeting in front, and separated on the vertex
by only a very narrow space. Antennae very short and broad, sordid yellow. Thorax
black and shining, almost destitute of pubescence ; the crenulations of the scutellum
very deep, some of them projecting behind as minute denticles. Squama and antisquama
white. Abdomen densely punctate, dull black, more shining at the base ; the basal
segment at the sides, with long, pale grey pubescence, on the dorsum with two small,
white, almost round marks : the second and third segments each with an elongate,
curved, white mark on each side. Legs black, extreme tips of femora and bases of
the tibiae sordid j'ellow. Hind leg with the basal joint of the tarsus very much enlarged,
and on the under side set with dense adpressed, pure white pubescence, which catches
the light in certain directions, and looks like quicksilver; the following joint also some-
what enlarged ; the under surface of the tibia also covered with white hair which is
not adpressed. One specimen.
ON THE INSECTS FEOM NEW BEITAIN. 391
Van der Wulp has figured the leg of E. argyropus (from New Guinea) in Termes.
Fiizetek, 1898, PI. XX. Fig. 6: in it the tarsus is less enlarged, and all the four
following joints are simple and symmetrical. E. argentipes Walk., according to the type
in the British Museum, has the hind feet very differently shaped.
Family ORTALIDAE. (Muscidae acalyptratae.)
Lamprogaster austeni, n. sp. PI. XXXV. Fig. 11 $ ; 11 «, 11 &, 11 c.
Thorace dorsoque abdominis viridi-purpureis, capite pedibusque flavis, illo vertice
fusco ; abdomine subtus niembranaceo utrinque versus apicem vesiculo protuberante ;
alis subopacis, basi et dimidio anteriore aurantiacis. Long. 12 m.m.
Antennae received in deep, elongate ear-like depressions ; first joint hardly visible,
second moderate, third elongate, twice as long as the second, bearing at its base an
arista twice as long as itself, and bare except for a few fine hairs at the base. Head yellow
with the vertex broadly fuscescent along the middle, a dark streak on each side below
the antenual cavity ; palpi yellow, labellum blackish. Thorax metallic, shining, feebly
pubescent, underface of scutellum yellow. Wings elongate, rendered dull by a dense dis-
tinct strigosity, the anterior part, and even the veins, dark yellow : squama ver}' large,
completely covering the halter. Legs clear yellow.
The metallic tint varies in colour, and may be in parts bluish or purplish.
The male has a large white, round vesicular prominence at each side of the abdomen
(Fig. 11 h), and the genitalia project between the pair of prominences. The female has
a very large, pap-like projection, instead of the round prominence of the male ; at the
tip, between the two paps, there is a rounded prominence from which projects the slender,
two-segmented ovipositor (Fig. 11 c).
The peculiar, vesicular structures of the abdomen shrivel after the insect is taken
from spirit and dried : and they also, I anticipate, only take on their full development in
life when the insect is sexually mature and occupied with reproduction. The species is
very like L. elonguta van der Wulp (from Batchian) but independent of the abdominal
structure it differs by its less elongate form, and by the colour of the undersurface of the
abdomen and hind coxae being yellow.
I have named this remarkable fly in honour of E. E. Austen, Esq., of the British
Museum, Natural History, who kindly assisted me in tracing its affinities. No one, un-
familiar with the intricate but unsatisfactory state of classification of Muscid flies would
dream of assigning a place to this insect amongst the Acalyptrate Muscidae, as the halters
are hooded in the most perfect manner.
GlRAFFOMYlA, n. g Muscidarum acalyptrataru
m.
Corpus elongatum, nitidum, pubescentiae destitutum. Prothorax elongatus ; caput
libcrum permobile, a thorace utrinque scleriti cervicale elongato separatum. Pectus valde
prominulum, ]ie(iibus intermediis et posterioribus contiguis, a pedibus anterioribus longe
remotis. Scutellum bispinosum. Caput marium interdum processu elongato ornatum.
53-2
392 ON THE IXSECTS FROM NEW BRITAIN.
This genus may be placed near Angitula Walker and Phytalmia Gerst. It is remark-
able on account of the elongation of the neck and its peculiar articulation ^vith the head
by means of an elongate cervical sclerite on each side ; and also by the prominence of the
breast whereby the middle legs are rendered contiguous with the posterior pair, and are
remote from the front pair. It also departs from Angitula by a slight peculiarity of the
wing-nervuration, the 3rd and 4th veins being more widely separated and the cross-vein
longer : in this respect it agrees with an insect from New Guinea, separated by Mr Austen
in the British Museum collection as a new genus allied to Angitula. The genus Phytalmia
Gerst. (Stett. Ent. Zeit. 1860, p. 169, PI. II. Fig. 3) has much in common with Giraffomyia
but in it the thorax is not elongated.
In some of the specimens the head is ornamented by a pair of peculiar large pro-
cesses that are apparently capable of movement by aid of a constriction placed near the
base. Several other Acalyptrate Muscids possess peculiar projections on the head. This
is well known in the case of the genus Elaphomyia. The genus Clitodoca also possesses
cephalic processes. In none of the forms is there however any trace of the projections
being divided into two segments except in the case of Giraffomyia. It is very curious if
it i^rove, as I expect it will, that so exceptional a structure should be present only in
some of the individuals of the male sex of the same species. Of six specimens the two
females do not possess the structure, and of the four males, one is entirely ^^^thout it,
while the other three possess it in very different degrees of development. When largely
developed it is accompanied by a considerable change in the form of the eye. The genus
Phytalmia has appendages somewhat like those of Giraffomyia, but not divided at the
base.
Walker places Angitula in the subfamily Sepsides. Gerstaecker has assigned no
position to Phytalmia.
Giraffomyia willeyi, n. sp. PI. XXXY. Fig. 12, ^ ; 12 a, profile of ^ ; 12 6, front of
head of male.
Gracilis, viridi-aenea, nitida, pedibus flavis, nigi'o-variegatis, alarum margine anteriore
argute nigi-a; capite aurantiaco, vertice fuscescente, ad marginem interiorem oculonim
albido-lineato. Long. 16 m.ni.
Mas ; capite utrinque processu magno, prope basin quasi articulato, apice laminam
curvatam flavam, nigi-o-marginatam formante.
Of this curious insect Dr Willey obtained at least two males. The colour of the
body and abdomen is a bras.sy-green, very shining. The head is pallid underneath the
antennae ; behind this it is of a tawny-orange colour, with a very fine white or silvery
line close to the eye : the vertex is fuscescent. The elongate cervical sclerites connecting
the head with the thorax are rather more than 1 m.m. long. The prothorax and abdomen
are membranous beneath. The surface is very polished and free from pubescence, but
there are punctures on the dorsal aspect of the thorax and some fine transverse wrinkles
on the me.sonotum. The long legs are yellow, with the tips of the femora and tibiae and
%vith the long tarsi blackish. The wings are transparent, with the front margin coloured
so as to form a very definite black stripe, and vnth this is connected a patch of black
ON THE INSECTS FROM NEW BRITAIN. ' 393
colour along the antero-external cross-vein. The only setae are, one behind each eye, and
a fringe of erect hairs along the free hind-margin of the vertex. The lialteres are white
and are placed at the junction of the perpendicular metanotum with the abdomen.
Although the specimens above alluded to are certainly males, they differ considerably
in the form of the peculiar cephalic appendages. Each appendage consists of two parts:
a basal portion, which is a direct continuation of the membranous undersurface of the
head, prolonged immediately beneath the eye and causing a large emargination therefore ;
and a second terminal part, which is harder and separated from the basal part bj' a deep
constriction ; at this point the appendage appears to be capable of a great deal of move-
ment. In one of the two specimens the basal prominence is very large ; while in the
other (PI. XXXV. Fig. 12, 12 a, 12 &) it is quite small.
Besides the two individuals described in the above lines, Dr Willey brought back
four other specimens considerably smaller in size, and concerning which I am in doubt as
to whether they may be a distinct species or small examples of G. willeyi. Two of them
are males, and two of them females. The smallest specimen is only 9 m.m. long. Both
the females and one of the males are entirely destitute of the peculiar cephalic append-
ages, while the other male possesses a vei-y small pair. I treat them as being a small
form of G. willeyi.
394 ON THE INSECTS FROM NEW BRITAIN.
EXPLANATION OF PLATE XXXV.
(Sharp, Insecta.)
Fig. L Profile of head and thorax of Xylotrupes gideon (1), male, from New Britain.
Fig. 2. Profile of head and thorax of Xylotrupes gideon male, from Java.
Fig. 3. Front of head and mandibles of various forms of Eurytrachelus intermedius from
New Britain ; viz. 3 a teleodont c? , 3 6 mesodout male, 3 c priodont male, 3 d female.
Fig. 4. Malacoderm Coleopterous larva ; 4 seen from above ; 4 a profile ; 4 b one of the
abdominal appendages more enlarged.
Fig. 5. Arrhenotus willeyi.
Fig. 6. Anterior legs of individuals of Xip/wtheata luctifera ; %-iz. 6 front leg of large
male ; 6 a front leg of small male ; 6 h front leg of female.
Fig. 7. Cacoschizus schmeltzi $.
jj ' ^. J) >j -f •
,, 7 h. „ „ hind foot of $ .
„ 8. Cacotrachelus sculptipennis S-
„ 8 a. „ ,, Profile of head and thorax of ? .
„ 9. Ithystenus debilis <? .
„ 10. Eurnems speculifer, hind leg.
„ 11. Lamprogaster austeni ?; 11a front of head; 116 uudersurface of abdomen of $ ;
lie undersurface of abdomen of female.
„ 12. Giraffomyia tmlltyi $ ; 12a profile; 12 6 front of head.
„ 13. Thynnus serriger $; 13 a extremity of abdomen seen from l.)ehind.
„ 14. Five cells from nest of Policies sp. illustrating the aberration of instinct described
on p. 388.
WiLLEY Zoological Results
Plate XXXV
Iff^ /^ ^
SHARP. INSECTA
Edwin WiUon C»ml)nel>jo
ON THE STOMATOPODA AND MACRURA BROUGHT BY
DR WILLEY FROM THE SOUTH SEAS.
By L. a. BORRADAILE, M.A.,
Lecturer at Selwyn College, Cambridge.
With Plates XXXVI.— XXXIX.
Dr Willey's collection of Stomatopoda and Macrura contains in all 82 species, of
which 20, rather less than a quarter, appear to be new to science. Notes on some of
these, short lists of references to others, and brief diagnoses of the new species in the
families Sqnillidae and Pontoniidae will be found in papers published by the Author
in the Proceedings of the Zoological Society ' and in the Annals and Magazine of Natural
History".
The specimens were collected in New Britain, the eastern archipelago of British
New Guinea, New Caledonia and the Loj'alty Islands, and the New Hebrides.
In New Britain the following forms were obtained : —
1. Protosquilla cerebralis Brooks [Pigeon Island].
2. Gonodactylus chiragra (Fabr.) [Talili Bay, Raluu].
var. anancyrus nov. [Talili Bay].
3. Odontodactylus scyllarus (Linn.).
4. Pseudosquilla ciliata (Fabr.) [Blanche Bay].
•5. Lysiosquilla biminiensis Bigelow, var. •pacificus nov. [Blanche Bay].
6. Penaeus fissurus Bate [Talili Bay].
7. Penaeus canaliculatus (Oliv.).
8. Stenopiis hispidus (Oliv.) [Blanche Bay].
9. Atya moluccensis de Haan [near Cape Gazelle].
10. Periclimenes parvus Borradaile [Blanche Bay].
11. Periclimenes tenuipes Borradaile [Ralun].
12. Pontonia ascidicola Borradaile [Blanche Bay].
13. Palaemo7i lar Fabr. [near Cape Gazelle].
14. Palaemon iveberi de Man [near Schultze Point].
15. Palaemonopsis willeyi sp. n. [Ralun].
1 P. Z. S., 1898, pp. 32, 457, and 1001. = A.nn. Mag. N. H. (7) ii. p. 377 (1898).
396 ON THE STOMATOPODA AND MACRURA BROUGHT BY
16. Pandalas {Parapandalus) serratifrons sp. n. [Blanche Bay].
17. Pandalus {Para'pandalus) tenuipes sp. n. [Blanche Baj'].
18. Pandalus (Parapandalus) longirostris sp. n. [Blanche Bay].
19. Heterocarpus ensifer A. M.-Edw. [Blanche Bay].
20. Synalpheus hiunguiculatus Stimps. [Blanche Bay].
21. Alpheus obesomamts Dana [Blanche Bay].
22. Alpheus laevis Randall [Blanche Bay].
23. Panulirus deniani nom. nov. [Blanche Bay].
24. Callianassa novae-britanniae sp. n.
25. Eiconaxius taliliensis sp. n. [Talili Bay].
26. Galathea elegans Adams and White.
27. Galathea grandirostris Stimps. [Talili Bay].
28. Munida scabra Henderson [Talili Bay].
var. longimanus nov. [Talili Bay].
29. Munida japonica Stimps. [Talili Bay].
30. Munida semoni Ortm. [Talili Bay].
31. Pagurus deformis H. M.-Edw. [Pigeon Isl.].
32. Pagurus asper de Haan [Blanche Bay].
33. Pagurus gemmatus H. M.-Edw.
34. Coenobita compressus H. M.-Edw. [Palakuvur].
35. Remipes admirabilis Thallw. [Blanche Bay].
36. Remipes ovalis A. M.-Edw.
37. Remipes celaeno de Man [Blanche Bay].
38. Albunea microps Miers [Blanche Bay].
In New Guinea: —
1. Periclinienes pai-asiticus Boiradaile [Milne Bay].
2. Anchistus miersi (de Man) [D'Entrecasteaux Group].
3. Anchistus biunguiculatus Borradaile [D'Entrecasteaux Group].
4. Conchodijtes meleagiinae Peters [Conflict Group. Engineer Group].
5. Pandalus {Parajxuidalus) tenuipes sp. n. [D'Entrecasteaux Group].
6. Saron marmoratus (Oliv.) [Louisiades].
7. Alpheus acjlaopheniae sp. n. [Engineer Group].
8. Callianidea tijpa H. M.-Edw. [Sariba].
9. Petrolisthes hastatus Stimps. [Sariba].
10. Petrolisthes lamarcki (Leach) [Sariba].
var. fimbriatus Borradaile [Sariba].
11. Pagurus deformis H. M.-Edw. [Conflict Group].
12. Pagurus setifer H. M.-Edw. [Conflict Group].
13. Pagurus euopsis Dana [Conflict Group].
In New Caledonia and the Loyalty Islands : —
1. Protosquilla cerebralis Brooks [Lifu].
2. Protosquilla trispinosa -Dana [Lifu].
DR WILLEY FROM THE SOUTH SEAS. 397
3. Gonodactylus chiragra (Fabr.) [Lifu].
var. smithi Pocock [Lifu].
var. anancyrus nov. [Lifu].
4. Squilla multituherculata Borradaile [Lifu].
5. Pseudosquilla ciliuta (Fabr.) [Uvea].
ti. Pericliiiienes spiniyerus (Ortin.) [Lifu].
7. Penclimenes lifaensis Borradaile [Lifu].
8. Goralliocaris inaequalis Ortm. [Lifu].
9. Leander pacijicus Stimps. [Isle of Pines].
10. Rhynchociiietes typiis H. M.-Edw. [Lifu].
11. Parhippolyte uveae sp. n. [Uvea].
12. Alpheinus tridens sp. n. [Lifu].
13. Synalpheus demani nom. nov. [Lifu].
14. Synalpheus neomeris (de Man) [Lifu].
15. Alpheus laevis Randall [Lifu].
16. Alpheus gracilidigitus Miers, var. [Isle of Pines].
17. Alpheus diadema Dana [Lifu].
18. Alpheus obesomanus Dana [Lifu].
19. Alpheus frontalis Say [Lifu].
20. Panulirus hispinosus sp. n. [Lifu].
21. Panulirus penicillatus (Oliv.) [Lifu].
22. Paribacus antarcticus (Rumph) [Lifu].
23. Scyllarus sieboldi de Haan [Lifu].
24. Galathea australiensis Stimps. [Lifu].
25. Galathea affinis Ortm. [Lifu].
26. Galathea spinimanus sp. n. [Lifu].
27. Porcellana sollasi Whitelegge [Lifu].
28. Petrolisthes hispinosus sp. n. [Lifu].
29. Petrolisthes lamarchi, var. fimbriatus Borradaile [Lifu].
30. Pachycheles sculptus (H. M.-Edw.) [Lifu].
var. tuberculatus nov. [Lifu].
31. Pachycheles lifuensis sp. n. [Lifu].
32. Pagurus asper de Haan [Lifu],
33. Pagurus deforrnis H. M.-Edw. [Lifu].
34. Pagurus setifer H. M.-Edw. [Lifu].
35. Pagurus gemmatus H. M.-Eihv. [Lifu].
36. Pagurus punctulatus Oliv. [Lifu].
37. Pagurus strigatus (Herbst) [Lifu].
38. Coenobita clypeatus (Herbst) [Lifu].
39. Coenobita perlatus H. M.-Edw. [Lifu].
40. Coenobita spinosus H. M.-Edw. [Lifu].
41. Coenobita coinpressus H. M.-Edw. [Lifu].
42. Coenobita rugosus H. M.-Edw. [Lifu].
43. Birgus latro (Linn.) [Lifu].
w. IV. 54
398 ON THE STOMATOPODA AND MACRURA BROUGHT BY
44. Bemipes testudinarius Latr. [Lifu].
45. Remipes pacificus Dana [Isle of Pines, Lifu].
46. Remipes celaeno de Man [Isle of Pines].
In the New Hebrides : —
Coenobita compressus H. M.-Edw.
The macmran fauna of the various localities, so far as it is revealed by the above
lists, will be seen to be essentially similar throughout. Further, with one exception
{Lysiosquilla biminiensis), all the known species are already recorded from the Indo-
pacific region.
It is interesting to notice that in several instances species on our roll are also
represented in the West Indies, either by forms so far as is known identical with
those from the Pacific, or by slightly different species or varieties.
The following is a list of such forms :
Indopacific form.
1. Gonodactylus chiragra (Fabr.)
2. Pseudosquilla ciliata (Fabr.)
3. Lysiosquilla biminiensis, var. pacificus nov.
4. Stenopus hispidus (Oliv.)
.5. Heterocarpixs ensifer A. M.-Edw.
6. Panulirus penicillatus (Oliv.)
7. Petrolisthes lamarcki (Leach)
West Indian representative.
G. oerstedi Hansen'.
P. ciliata, var. occidentalis nom. nov.^
L. biminiensis Bigelow.
S. hispidus.
H. ensiferr.
P. guttatus (Fabr.)l
P. lamarcki, var. asiaticus (Leach)*.
The following species in Dr Willey's collection occur also in sub-tropical
Australia : —
1. Protosquilla trispinosa (Dana) [Swan River']
2. Gonodactylus chiragra (Fabr.) [Swan River'].
3. Penaeus canalicidatus (Oliv.) [Port Jackson => "].
4. Alpheus laevis Randall [Port Jackson'].
5. Galathea australiensis Stimps. [Port Jackson''*].
6. Pagurus deformis H. M.-Edw. [Port Jackson"].
7. Coenobita rugosus H. M.-Edw. [Sydney'].
8. ? Coenobita spinosus H. M.-Edw. [Sydney'].
The following in New Zealand": — •
1. Protosquilla trispinosa (Dana).
2. Palaemon lar Fabr.
3. Rhynchocinetes typus H. M.-Edw.
1 See below, under G. chirnijra. ^ See below, under P. ciliata.
- Ortmann, Zool. Jahrb. vi. Syst. p. 29. ■* See P. Z. S., 189S, pp. 464 — 467. Var. asiaticus is also Indopacific.
5 Haswell, Cat. Austral. Crust., Syduey 1882. « Whitelegge, J. E. Soc. N.S.W. xxiii. pp. 224, 232.
' Miers, Cat. N. Z. Crust, pp. 77, 87, 90.
DR WILLEY FROM THE SOUTH SEAS, 399
Turning now to a systematic survey of the material, we may notice at the outset
the relative extent to which the various groups of Macrura and Stomatopoda are
represented.
There are seven species of Stomatopoda, belonging to six out of the nine genera
of that order. Of these species one, a Squilla, is new. lu the Macrura. the Penaeidea
are represented by two species of Penaexis, the Stenopidea by one species of Stenopus,
the Caridea by 32 species from 17 genera, the Loricata by .5 species from 3
genera, the Thalassinidea by three species from as many genera, and the Anomala bv
32 species from 9 genera. The 19 new species in this suborder are distributed
as follows: in the Caridea there are 13, in the Thalassinidea 2, in the Loricata one,
and in the Anomala three.
Of the 38 genera in the whole collection, the most numerously represented is
Alpheus, with six species ; PericUinenes, on the other hand, has both absolutely and
relatively' the largest number of new species (four out of five).
Conspicuous by their absence are the Astacidea and the Crangoninea.
Sub-class. THORACOSTRACA.
Order. STOMATOPODA.
Family. Squillidae.
Genus. Protosquilla, Brooks, 1886.
1. Protosquilla cerebralis Brooks, 1886.
Protosquilla cerebralis Brooks, 'Challenger' Stomatopoda, p. 72, PI. XIV. Figs. 2
and 3, XVI. Figs. 2 and 3 (1886); Borradaile, P. Z. S. 1898, PL V. Fig. 6a.
Brooks's specimens of P. cerebralis were all females. Fortunately, however, Dr
Willey's collection contains a male specimen, so that it has been possible [P. Z. S.
loc. cit.l to figure for this species the peculiar structure on the endopodite of the fir.«t
abdominal appendage of male Stomatopoda.
The outer leaf of the last joint of this organ is smaller than the inner, <.ii
which it is borne as a lobe, and from which it is not sundered by a suture. The
fixed limb of the pincers is hook-shaped, while the movable limb is bent sharply
and of a shape something like that of a boomerang. The first joint is produced on
the outside at the free end.
Two % from Sandal Bay, Lifu, Loyalty Islands. One ^ from Pigeon Island, New
Britain.
' With the exception, of course, of the new genera Palaemonopsis, Parhippolyte, and Alpheinus, each
founded for a single new species.
54-2
400 ox THE STOMATOPODA AND MACRURA BROUGHT BY
2. Protosquilla trispinosa (Dana), 1852.
Gonodactylus trispinosus, White, List Crust. Brit. Mus. p. 75 (1847) nom. nud. ;
Dana, Zool. U. S. Expl. Espd., Crust, i. p. 623 (1852); Miers, Cat. N. Zeal. Crust.,
p. 90; Ann. Mag. X. H. (5) v. p. 121, PL III. Fig. 10 (1880); Haswell, Cat. Austral.
Crust., p. 211 (1882).
Protosquilla trispinosa Brooks, Challenger Stomatopoda, p. 71 (1886); Borradaile,
P. Z. S., 1898, p. 34, PI. V. Figs. 1, la (1898).
The following is an amended diagnosis of this species : —
"A Protosquilla with the lateral spines of the rostrum nearly as long as the
median; carapace with angles nearly rectangular, anterior more acute than posterior;
fifth and eighth thoracic segments with the lateral margin subacute, sixth and seventh
with the same somewhat square ; first abdominal segment with one, second to fourth
with two lateral sulci ; fifth longitudinally corrugated ; sixth with six tubercles bearing
fine spinules, and clearly marked off from the telson ; the latter with a median and
two lateral large tubercles covered with fine spinules, the median anterior to the two
laterals, and with the posterior border divided by deep narrow fissures into six lobes ;
submedian, intermediate and lateral spines of the telson small and sunk in notches,
several submedian spinules; outer spine of basal prolongation of uropod (6th abdominal
limb) larger than inner, not armed with a tooth on its inner margin."
Length about 40 mm.
For figures see P. Z. S. 1898, PI. V. Figs. 1, la.
1 $ from Lifu, Loyalty Islands.
Genus. Gonodactylus Latr., 1825.
3. Gonodactylus chiragra (Fabr.), 1793.
Squilla chiragra, Fabricius, Ent. Syst. in., 1, p. 513 (1793).
Gonodactylus chiragra Latreille, Encycl. Meth., x. p. 473 (1825) ; Miers, Ann.
Mag. N. H. (5) V. p. 118 (1880); Haswell, Cat. Austral. Crust, p. 210 (1882);
de Man, Zool. Jahrb. x. Syst. p. 694, PL XXXTIII. Fig. 77 (1898); Borradaile, P. Z. S.
1898, p. 34, Pis. V. Fig. 4, and YL Fig. 8.
Gonodactylus smithii Pocock, Ann. Mag. N. H. (6) xi. p. 475, PL XX. B, Fig. 1 (1893).
De Man (loc. cit.) selects as the type of this species the form with the middle
keel of the telson anchor-shaped. He further separates from it a variety which he
names acutirostris and characterises as follows : —
1. Middle keel of telson not anchor-shaped.
2. Keels of submedian spines of telson converge from behind forwards.
3. Outer angles of rostrum acute.
There are in the present collection four specimens of a variety intermediate between
the type and var. acutirostris. .In these specimens the telson has not the anchor-
DR WILLEY FROM THE SOUTH SEAS. 401
shaped middle keel ; but neither are the outer angles of the rostrum sharp, and the
keels of the sabmedian telson-spines do not converge forwards. Should a name for
this variety be thought necessary, it is here proposed to call it var. anancyrus.
To these three varieties may be added that distinguished by Pocock in 1893
{loc. cit.) under the name of G. smithii, and characterised as follows': —
1. The keels of the sixth abdominal segment and telson are more compressed
than in the type.
2. The keels of the sixth abdominal segment are produced, without constriction
into long spines.
3. The upper edge of the middle keel of the telson is almost straight, and is
produced backwards into a spine.
4. The 'flukes' of the anchor on the telson are represented by two narrow ridges
running forwards from the hind end of the middle keel.
5. On each side of the first five abdominal tergites is a small, sharply-defined, dark spot.
Lastly, the Gonodactylus oerstedi of Hansen^ may be considered in connection with
the above forms, since it differs from them no more than they from one another,
and rests its claim to specific rank mainly on its geograiihical distribution. It is
characterised by the presence of a small swollen ridge on the inside of the keel of
the intermediate spine of the telson, and in other respects resembles the type variety
of G. chiragra. It is at present known only from the West Indies and east coast of
America.
The distinguishing marks of the above forms may be set forth in key form as
follows : —
1. Without a swollen ridge on the inside of the keel of the intermediate telson-
spine. Distribution Indopacific.
2. Keels of sixth abdom. segment and telson rounded, not produced without
constriction into spines. Middle keel of telson without a spine.
3. Middle keel of telson anchor-shaped.
Variety A. (type).
3'. Middle keel of telson not anchor-shaped.
4. Outer angles of rostrum not acute. Keels of submedian telson-spines not con-
verging forwards.
Variety B. {anancyrus).
4'. Outer angles of rostrum acutely pointed. Keels of submedian telson-spines
converging forwai'ds.
Variety C. (acutirostris).
' P. Z. S., 1893, p. 34, where it is claimed that this form is but a variety of G. chiragra.
2 Plankton-exped., laop. Cum. u. Stem., p. 65. See also P. Z. S. loc. cit., PI. V., Fig. 3.
402 ON THE STOMATOPODA AND MACRURA BROUGHT BY
2'. Keels of sixth abdom. segment and telson compressed. Keels of sixth abdom.
segment produced without constriction into long spines. Middle keel of telson ending
iti a sjDine.
Variety D. {smithi).
V. With a swollen ridge oa the inside of the keel of the intei'mediate telson-
spine. Distribution Atlantic.
Variety (?) E. {G. oerstedi Hansen).
The specimens in the present collection include : —
i. Var. A. (type). 1 J' and 1 $ from the Isle of Pines, New Caledonia ; 2 J
and 3 $ from Lifu, Loyalty Islands; 1 ? from Talili Bay, New Britain; 1 $ from
Ealun, New Britain.
ii. Var. B. (anancyrns). 1 j/ from Talili Bay ; 2 $ from Lifu ; 1 j/ loc. ?
iii. Var. D. (smithi). 2 j" and 2 $ from Lifu.
Genus. Odontodactylus Bigelow, 1895.
4. Odontodactylus scyllarus (Linn.), 1758.
For references see P. Z. S. 1898, p. 36.
1 % from New Britain.
Genus. Pseudosquilla Dana, 1852.
5. Pseudosquilla ciliata (Fabr.), 1793.
Squilla ciliata, Fabricius, Ent. Syst. ill., 1, p. 512 (1793).
Pseudosquilla ciliata, Miers, Ann. Mag. N. H. (5) V. p. 108 (1880); Brooks,
'Challenger' Stomatopoda, p. 53, PI. XV. Fig. 10 (1886); Borradaile, P. Z. S. 1898^
p. 36.
The present specimens agree with that of Brooks from Honolulu and that brought
by Mr Gardiner from Funafuti (P. Z. S. loc. cit.) in the following points in which
they differ from Brooks's West Indian specimens.
1. The fourth abdominal segment has no spine at the hinder angle.
2. The inner spine of the basal prolongation of the uropod is longer than the
outer.
Should these differences prove to be characteristic of the forms from the two
regions the name of var. occidentalis would be a suitable one to apply to that from
the West Indies.
1 % from Uvea, Loyalty Islands. 1 ^ from Blanche Bay, New Britain.
DR WILLEY FROM THE SOUTH SEAS. 403
Genus. Lysiosquilla Dana, 1852.
6. Lysiosquilla bimimensis Bigelow, 1893, var. pacificus uov.
Lysiosquilla himinieTisis Bigelow, Job. Hop. Univ. Circ. c\'l., p. 102 (1893); Proc.
U. S. Nat. Mus. XVII. p. 504, Figs. 4—7 (1895).
A single male specimen from New Britain seems to belong to a variety of this
species. The resemblance to Bigelow's figures and description is complete save in the
following small points : —
1. The movable submedian spines of the telson are stouter than in Bigelow's
figure, while the innermost pair of submedian spinules are minute.
2. The outer spine oa the basal prolongation of the uropod is slightly longer
than it is figured for the type.
3. The antennal scales are somewhat smaller.
■4. In addition to the markings shown by Bigelow there is a narrow band of
deep black (in spirit) on the hinder edge of each segment from the sixth thoracic to
the fifth abdominal inclusive.
These differences do not seem sufficient to justify the separation of the form in
question from the West Indian species, and it is accordingly proposed to call it var.
pacificus, emphasizing thereby its interesting distribution. L. bimimensis is the only
species in the collection already known and not recorded from the Indopacific region,
and forms one of the list already given to illustrate the coincidences between the
Macruran and Stomatopodan fauna of the West Indies and that of the South Sea
Islands visited by Dr Willey.
Genus. Squilla Fabr., 1793.
7. Squilla multituberculata Borradaile, 1898.
Squilla multituberculata, Borradaile, P. Z. S. 1898 p. 38, PI. VI. Fig. 7, 7a — 7c.
The short diagnosis accompanying the figures of this species in the above-mentioned
paper may be here amplified with certain further details.
The rostrum is sub-rectangular, somewhat narrower behind than before, without
carinae, and with the antero-lateral angles produced, bent downwards, and acute.
The eyes are elongate and flattened from above downwards, and their cornea
consists of two roughly hemispherical portions set side by side on the end of the stalk.
The carapace is small, narrower before than behind, with rounded angles and the
hinder border somewhat concave. The sides of the fifth thoracic segment are sharp,
those of the sixth to eighth subtruncate.
The antennae have long stalks, and the last joint of the scale of the second pair
is small.
In the great claw the last joint is stout, bearing on the inside four teeth (in-
cluding the terminal tooth), and on the outside three short teeth at the base.
The tail fin. The sixth abdominal segment bears eight roughly longitudinal ridges
and a few scattered knobs. It has two small blunt processes on the hinder edge, and
404 ON THE STOMATOPODA AND MACRURA BROUGHT BY
is sharply marked oflf from the telson. The latter is rather strongly convex, and is
covered with small blunt spines. Along the middle line runs a raised ridge, grooved
above. The marginal spines are small, and can hardly be seen from above. The sub-
medians have a movable tip. There are four or five submedian spinules and five or
six lateral. The uropod has a large basal joint, with the outer of the two spines on
its prolongation obsolescent. The two joints of the exopodite are subequal, the first
bearing eight spines on the outside. The endopodite is shorter than the exopodite.
The latter equals the telson.
In the male the basal joint of the endopodite of the first abdominal appendage
is broad and armed with long stout hairs. The inner leaf of the end-joint is some-
what narrow. The outer leaf is narrow, and is shorter than the inner, behind which
it is hidden ^ The limbs of the pincers are long and narrow.
1 (/ and 2 %, from Sandal Bay, Lifu, Loyalty Islands.
Order. DECAPODA.
Sub-Order. MACRURA.
Tribe. PENAEIDEA.
Family. Penaeidae.
GE^X'S. Penaeus Fabr., 1798.
8. Penaeus fi^suriis Bate, 1888.
Penaeus fissurus Bate, " Challenger," Macrura, p. 2G3, PL XXXVI. , Fig. 1.
1 $ from Talili Bay, New Britain.
9. Penaeus canaliculatus (Oliv.), ISll.
Palaemon canaliculatus, Olivier, Encycl. Meth. viii. p. 660 (1811).
Penaeus canaliculatus, H. M. -Edwards, H. X. Crust. II. p. 414 (1837) ; Bate
"Challenger," Macrura, p. 243, PL XXXII. Figs. 1, 2 (1888).
1 (/ fi-om New Britain.
Tribe. STENOPIDEA.
Family. Stenopidae.
Genus. Stenopus Latr., 182.5.
10. Stenopus hispidus (Oliv.), 1811.
Palaemon hispidus, Olivier, Encycl. Meth. viii. p. 666, PL XIX., Fig. 2 (1811).
Stenopus hispidus, Latreille, Desmarest's " Consid. s. 1. Crust.," p. 227 (182.5) ; Adams,
Voy. ' Samarang,' p. 61 (18.50) ; Herrick, Mem. Nat. Ac. Sci. v. 4, p. 348, Pis. V., XIII.
1 Thus it is not shown in the figure gi%'eu in the P. Z. S. loc. cit. The lobe underlying the movable
limb of the pincers in this figure was drawn in error and does not exist.
DR WILLEY FROM THE SOUTH SEAS. 405
The specimens agree with Herrick's description of the West Indian examples com-
pletely, even in the points in which the latter differ from Adams's figures.
2 c/ and 1 $ from Blanche Bay, New Britain.
Tribe. CARIDEA.
Family. Atyidae.
Gends. Atya Leach, 1S17.
11. Atya moluccensis de Haan, 1849.
Atija moluccensis, de Haan, Faun. Japon., Crust., p. 186 (1849) ; Miers, Ann. Mag.
N. H. (.5) V. p. 382, PI. XV., Figs. 3, 4 (1880); de Man, in Max Weber's "Zool. Ergebnisse,"
u. p. 357, PI. XX., Fig. 20 (1892).
1 (/ and 1 ? taken in a stream near Cape Gazelle, New Britain.
Family. Pontoniidae.
Further particulars are now added to the j)reliminary diagnoses of certain species in
this family already published b}' the author in the " Annals and Magazine of Natural
History," 1898.
Genus. Periclimenes Costa, 1844.
Periclimenes, Costa, Ann. Ac. Aspir. Nat. Nap. Ii. (1844); Faun. Regn. Nap. II.
1 (1846); Borradaile, Ann. Mag. N. H. (7) n. p. 380 (1898).
Pelias, Roux, Mem. s. 1. Salicoques, p. 2n (1831) nom. praeoc.
Anchistia, Dana, U. S. Expl. Expd. Crust. I. p. 577 (1852).
Bennisia, Norman, Ann. Mag. N. H. (3) VIU. p. 278 (1861).
The species described as Pelias migratorius by Heller in 1862 was afterwards
placed by the same author in his new genus Palaemonetes, and recognised as identical
■with P. varians (Leach). It is therefore erroneously placed in the genus Periclimenes
in the revision of that genus by the present writer (Ann. Mag. loc. cit.).
12. Periclimenes spinigerus (Ortm.), 1890.
Anchistia spinigera, Ortmann, Zool. Jahrb. v. Syst., 3, p. 511, PI. XXXVI., Figs. 23,
23 a (1890).
Periclimenes spinigerus, Borradaile, Ann. Mag. N. H. (7) II. p. 383 (1898).
1 $ from Lifu, Loyalty Islands.
13. Periclimenes lifuensis Borradaile, 1898, Figs, la — Ic'.
Periclimenes lifuensis, Borradaile, Ann. Mag. N. H. (7) ii. p. 384 (1898).
Carapace. The rostrum is straight, outreaches the antennal scale, and bears six
teeth above but none below. The supraorbital and antennal spines are present, but
not the hepatic. There is also a spine on the mid-dorsal line behind the rostrum.
The pterygostomial angle is acute.
1 This reference (and all similarly placed references) relates to the figures on Plates XXXVI. — XXXIX., the
numbers of which run consecutively.
w. IV. 55
406 ON THE STOMATOPODA AXD MACRURA BROUGHT BY
Antennae. The stalk of the first antenna is shorter than the scale of the second,
and consists of a very broad first joint, projecting forwards and outwards in a sharp
point at the distal end, and two short, subequal following joints. The inner fiagellum
is broken short on either side in the present specimen, and the thicker part of the
outer outreaches the scale of the second antenna, though not the fringe of hairs on
that structure. The stalk of the second antenna about equals that of the first in length.
The thiJ-d nuuvilliped reaches the end of the stalk of the second antenna.
Legs {Pereiopoda). The first pair of legs has the beginning of its wTist-joint
(carpopodite) even with the end of the second joint of the antennular stalk, and
outreaches the scale of the second antenna by the hand (propodite) and the last
two-thirds of the -svi-ist. The fingers about equal the palm in length. The second pair
are subequal. Their meropodite about reaches the end of the rostrum, the wrist is
short and armed above with a spine, and the fingers are about two-thirds as long
as the palm, hairy, and curved towards one another at the tip, thus enclosing a space.
The remaining legs are short and stout, and their last joint is curved and arises among
a tuft of long hairs.
Tail-fin. The uropods are longer than the telson, and their exopodite and endo-
podite are subequal.
Length of the single specimen 11 mm.
From the above characters it would seem to be necessary to place this species
in the neighbourhood of P. gracilis (Dana) 1852, from which, however, it is sundered
by the presence of a supraorbital spine, to mention only one point of difference.
Lifu, Loyalty Islands.
14. Periclimenes tenuipes Borradaile, 1898, Figs. 2a — 2/
Periclimenes tenuipes, Borradaile, Ann. Mag. N. H. (7) ii. p. 384 (1898).
Carapace. The rostrum is long, slender, curved upwards, armed above with ten
teeth (of which the first two stand on the carapace) and below with seven, and out-
reaches the antennal scale, but not the outer antennular flagellum. Hepatic and
antennal spines are present, and the pterygostomial angle is rounded.
All the appendages are unusually slender and elongated.
Antennae. The stalk of the first antenna is shorter than the scale of the second.
Both its flagella are long, the outer being bifid at the tip. The stalk of the second
antenna does not reach the end of the first joint of that of the first. Its scale is
longer than the antennular stalk, shorter than the rostrum, and narrow.
Mouth-limbs. These are shown in Figs. 2c—;/'. The third masilliped nearly reaches
the end of the first joint of the antennular stalk.
Legs. The first pair is wanting in the single specimen. The end of the meropodite
of the second is even with the rostrum, and is armed beneath with a spine. The wrist
is longer than the meropodite and slightly longer than the palm. It grows broader towards
its outer end, where it is armed above with a spine. The movable finger bears three
teeth on its inner edge. In the last three legs the propodite is armed with spines.
Tail-fin. The exopodite of the uropod is longer than the endopodite, and both
are considerabh' longer than the telson. The latter structure is armed at its free end
DR WILLEY FROM THE SOUTH SEAS. 407
with the usual six spines found in this position in the present family. These are,
namely, two subniedian, two intermediates longer than the submedian, and two laterals
shorter than the submedian.
The single specimen measures 11 mm. in length, and was found on the reef at
Ralun, New Britain.
The following two species are both of small size and have a certain immature
appearance. They are here described and named provisionally.
15. Periclimenes parvus Borradaile, 1898, Figs. 3a — 3c.
Periclimenes parvus, Borradaile, Ann. Mag. N. H. (7) il. p. 384 (1898).
Carapace. Rostrum slightly longer than antennular stalk, bent downwards at first,
but tending to straighten towards the tip, above with a deep crest bearing six teeth,
below with one tooth. Antennal and hepatic spines are present, and the pterygostomial
angle is subrectangular.
Antennae. The stalk of the first antenna is shorter than the scale of the second.
The stalk of the second antenna does not reach the end of the first joint of the
antennular stalk ; the scale is outreached by the antennular flagella.
The eyes are large.
The third maxilliped is shown in Fig. 3c; it is rather small, barely reaching the
end of the peduncle of the second antenna.
The legs. The first pair of legs are short, not outreaching the antennal scale,
and fairly stout. The second pair are short, simple in form, without spines, and out-
reach the antennal scale by about the latter half of the meropodite. The longest joint
is the hand. The remaining legs are slender, and have nearly straight, biuuguiculate
dactyles.
The tail-fin. The exopodite of the uropod is longer than the endopodite, and both
are longer than the telson. The latter bears at its hind end two very strong spines
and four weaker ones.
Two specimens, 8".5 mm. long, were taken at Rakaiya, Blanche Bay, New Britain.
16. Periclimenes parasiticus Borradaile, 1898, Figs. 4a — 46.
Periclimenes parasiticus, Borradaile, Ann. Mag. N. H. (7) ii. p. 384 (1898).
Carapace. The rostrum is straight, with a large convex donsal crest of seven
teeth, but unarmed below. It just outreaches the first joint of the antennular stalk.
Antennal spines are present, but not supraorbital.
The eyes are large.
Antennae. The stalk of the first antenna is shorter than the scale of the second.
The stalk of the second is not so long as the basal joint of the first. The scale
of the second antenna is broad and reaches the end of the thicker antennular tlagellum.
The third maxilliped barely reaches the pter3-gostomial angle of the carapace.
Legs. First pair short, strong, and with unusually stout hands. Second pair small
with short wrist, and without spines on any of the joints.
The sixth abdominal segment is considerably elongated.
55—2
408 ON THE STOMATOPODA AXD MACRURA BROUGHT BY
Tail-fin. The exopodite of the iiropod is longer than the endopodite. This, in
turn, is longer than the telson.
Length of largest specimen 7 mm.
Four specimens of this species were found living among the spines on the back
of a black starfish of the genus Linckia.
Genus. Anclnstus, Borradaile, 1898.
17. Anchistus miersi (de Man), 1888.
Harpilius Miersi, de Man, Journ. Linn. See, Zool. xxii. p. 274, PL XXII. Figs.
6—10 (1888).
Anchistus miersi, Borradaile, Ann. Mag. N. H. (7) ii. p. 387 (1898).
2 $ and 1 </, found in the mantle-chamber of a Tridacna squamosa at Dobu,
D'Entrecasteaux Group, British New Guinea.
18. Anchistus biunguiculatvs Borradaile, 1898, Figs, oa — 5c.
Anchistus hiunguiculatus, Borradaile, Ann. Mag. N. H. (7), ii. p. 387 (1898).
Carapace. The rostrum reaches the end of the first joint of the antennular stalk,
is strongly curved downwards, and bears no teeth. The antennal spine alone is present,
and the pterygostomial angle is rounded.
Antennae. The stalk of the first antenna does not quite reach the end of the
scale of the second. The flagella are subequal and of moderate length onl}'. The
stalk of the second antenna is as long as the basal joint of the first, and the scale
is broad.
Mouth-limbs. These have not been examined in the single specimen. The third
maxilliped reaches half way up the last joint of the second antennal stalk.
Legs. The first pair of legs outreach the antennal scale by the last half of the
wrist and the hand. The second pair are equal, symmetrical, and without spines on
any of their joints. The hand is long and rather narrow. The wrist is half the
length of the palm. The fingers are unequal, the movable one being considerably
longer than the immovable and hooked at the end. The immovable finger is more
than half as long as the palm.
Tail-fin. The exopodite of the uropod is slightly longer than the endopodite.
The latter is somewhat longer than the telson, which is triangular, with a rounded
apex bearing the usual six spines.
Length .50 mm.
One ? fi'om a Tridacna, in Tubetube, Engineer Group, British New Guinea.
Genus. Coralliocaris Stimpson, 1860.
19. Coralliocaris inaequalis, Ortmann, 1890.
Coralliocaris inaequalis, Ortmann, Zool. Jahrb. V. syst. 3, p. 510, PI. XXXVI.
Figs. 21, 2ld—i (1890).
3 specimens from Sandal Bay, Lifu, Loyalty Islands.
DR WILLEY FROM THE SOUTH SEAS. 409
Genus. Pontania Latr. 1829.
20. Fontonia ascidicola Borradaile, 1898, Figs. 6ct — 66.
Pontonia ascidicola, Borradaile, Ann. Mag. N. H. (7) ii. p. 389 (1898).
Carapace. Rostrum short, reaching only half way up the first joint of the
antennular stalk. The free end is strongly curved downwards and lacks an inferior
keel. The antennal spine alone is present and the pterygostomial angle is rounded.
Antennae. The stalks of the two pairs of antennae and the scale of the second
are subequal. The scale is broad and of but moderate length.
Mouth-limbs. These have not been dissected out. The third maxillijjud ends opposite
the second joint of the antennular stalk (Fig. 66).
Legs. The first pair of legs are rather unusually strong. Then- meropodite reaches
the end of the first joint of the antennular stalk. The wrist is a little shorter than
the meropodite and the hand a little shorter than the wrist. The fingers are as long
as the palm. The second pair are unequal. In the larger, the hand is of great size, the
fingers half the length of the palm, the wrist short and stout, the meropodite longer than
the wrist, the movable finger bearing one tooth and the immovable finger two. In
the smaller leg of the second pair, the hand is still the longest joint, the movable
finger is narrow and crosses its fellow at the tip, and both are provided with teeth
as in the longer hand. In the female both legs of the second pair are relatively
shorter than in the male. The dactyles of the remaining legs are short, fairly stout,
and provided with several spines underneath.
The abdominal pleurae are greatly developed in the female.
Tail-fin. The two rami of the uropod are subequal in the male. In the female
the endopodite is the longer. In each case the telson is as long as the exopodite,
and bears the usual six spines at the free end.
The length is 13 mm.
1 (/• and 1 ? from Blanche Bay, New Britain.
Genus. Conchodytes Peters, 18.51.
21. Conchodytes meleagrinae Peters, 18.51.
For references see P. Z. S. 1898, p. 1007.
3 ?, from Sandal Bay, Lifu, Loyalty Islands, and from Engineer Group and Conflict
Group, British New Guinea, respectively.
Family. Palaemonidae.
Genus. Palaemon Fabr., 1798.
22. Palaemon lar Fabr., 1798.
For references see P. Z. S. 1898, p. 1008.
4 t/" of various ages, taken near Cape Gazelle, New Britain.
410 ox THE STOMATOPODA AND MACRUKA BROUGHT BY
23. Pulaemon weberi de Man, 1892.
Palaemon webe7-i, de Man, in Max Weber's " Zool. Ergebnisse.," II. p. 421, PI. XXV.
Fig. 23 (1892).
One young male (.5-5 mm.), agreeing closely with de Man's description of a similar
specimen from the East Indies, was taken in a stream near Schultze Point, New
Britain. In both chelae, however, the fingers are shorter than the palm, while the
whole body is smooth, neither carapace nor telson being " kornig rauh."
Genus. Leander Desmarest, 1840.
24. Leander pacificus Stimpson, 1860.
Leander pacificus, Stimpson, Proc. Ac. N. Sci. Philad. 1860, p. 40.
The iifth pair of legs in the single specimen seem somewhat longer than is
indicated by Stimpson's description.
Locality, Isle of Pines.
Genus. Palaemonopsis nov.
There is in the collection a solitary Palaemonid for which it seems to be
necessary to found a new genus. The specimen in question differs from the members
of the genus Palaemon in the absence of a mandibular palp. From Palaemonetes it
differs in having on each side of the carapace one antennal spine only, and, directly
behind the eye, at a short distance from the edge of the carapace, a large, blunt,
roughly triangular process. About half of the thicker branch of the outer flagellum
of the first antenna is fused with the thinner branch, but the two branches are quite
distinctly formed down to their bases, so that the genus must be placed in the
present family rather than in the Pontoniidae. The slenderness of the third maxilliped
points to the same conclusion.
25. Palaemonopsis willeyi sp. u., PL XXXVI. — XXXVII., Figs. 7a — 7e.
Diagnosis : — " A Palaemonopsis with the rostrum straight, bearing six equal teeth
above and four teeth below, outreaching the antennular stalk but not the antennal
scale ; carapace bearing a single antennal spine on each side, and a large triangular
process behind the eye ; pterygostomial angle subrectangular ; first antenna with last
two joints of the stalk together shorter than first joint, and subequal ; flagella unequal,
the outer larger and with its two branches fused for about half the length of the
thicker branch ; second antenna with the stalk equal to the first two joints of the
antennular stalk, the scale longer than the rostrum, narrowing to the free end, which
is truncated and bears a triangular tooth, projecting beyond it, on the outside ; third
maxilliped small and slender; first pair of legs reaching the end of the antennal scale,
with wrist and meropodite subequal and longer than the hand ; second pair large, strong,
longer by the hand than the antennal scale, with short, stout wi-ist, and meropodite a little
longer than the palm, the fingers longer than the palm, crossing at the tip and serrate,
none of the joints with spines ; remaining legs fairly stout, with small, straight, slender
DR WILLEY FROM THE SOUTH SEAS. 411
dactyles, third longer than fourth or fifth ; sixth abdominal tergite with a broad tri-
angular median backward projection, flanked on each side by a spine ; endopodite of the
uropod very slightly longer than the telson, exopodite slightly larger than endopodite ;
telson elongate, narrowing gently towards the free end, which is truncate, bearing on
each side a short, strong spine, and in the middle a tuft of hairs, dorsal surface with
four pairs of movable spines."
Length 30 mm.
1 specimen from Ralun, New Britain.
Family. Pandalid.\e.
Genus. Pandalus Leach, 1814.
Into this genus Ortmann has reunited the genera Plesionika, Nothocaris, and
Pandalopsis of Bate. Three species of Pandalidae from New Britain have certain
characters in common which appear to justify the foundation for them of a new sub-
genus equivalent to the above-mentioned groups of species.
Subgenus. Parapandalus nov.
Characters : —
1. Carapace without lateral carinae.
2. Rostrum long, slender, armed above and below with movable spines.
3. First antenna with long flagella and pointed stylocerite.
4. Hinder lobe of scaphognathite truncated.
5. Third maxilliped with an exopodite.
6. Fii-st leg subchelate' owing to a small projection of the propodite at the base
of the finger.
7. Second pair of legs equal, with 25 — 30 joints in the wrist.
8. Eye with large cornea, well-marked ocellus, and two-jointed stalk.
9. Gill formula as in Pandalus (sens, str.), save that in two of the species
epipodites are wanting behind the third maxilliped.
26. Pandalus (Parapandalus) serratifrons sp. n., Figs. 8« — Sd.
Diagnosis : — " A Pandalus with the rostrum long, outreaching the antennal scale,
armed above and below with numerous small, similar, movable .spines, of which the
1 The word subehelate hardly describes the structures in question satisfactorily. The impresBion
is not that of a practicable grasping organ. Reference to fig. M will make this clear. For some interesting
i-emarks on the subject of this limb in Pandalus see Caiman, Ann. Mag. N. H. (7) in. p. 27 (1899).
412 ox THE STOMATOPODA AND MACRURA BROUGHT BY
first four or five are on the carapace ; carapace with antennal spine, acute pterygo-
stomial angle, and a small dorsal carina on the anterior third of its length; eyes
fairly large, with ocellus distinct, but not completely separated fi-om main cornea; all
the appendages long and slender ; first antenna with relatively short stalk, first joint
longer than second and third together, third longer than second, both flagella long, outer
broad and flattened at the base and bearing in this region a fi-inge ; stylocerit eas long
as first joint and ending in a sharp point ; second antenna with the stalk short,
equal to the first joint of the antennular stalk; scale elongate, narrowing to the
free end, where it is truncated, with firm outer edge ending in a spine which
starts before the end of the scale and projects beyond it ; third masilliped longer
by its last two joints than the antennal scale ; epipodites wanting behind the third
maxilliped ; legs of the first pair longer by their last two joints than the third
maxilliped ; legs of the second pair equal, exceeding the antennal scale by the hand
and the last two or three joints of the wrist, the wrist about 25-jointed, with the
last joint nearly equal to three of the preceding joints, growing broader towards the
hand; hand arising in a tuft of hairs and with long hairs on the fingers; remaining
legs long and slender, with movable spines on the meropodite and the dactyle small ;
uropod with exopodite larger than endopodite, endopodite longer than telson ; telson
lono-, very narrow, with four spines at the end, and four pairs of spines on the dorsal
surface."
Length of largest specimen 85 mm.
This species forms part of the food of the Nautilus.
7 J", trawled at depths of 50 — 100 fathoms in Blanche Bay, New Britain.
27. Pandalus (Parapandalus) tmidpes sp. n.. Fig. 9.
Diagnosis : — " A Pandalus with the rostrum long, outreaching the antennal scale,
bent slightly upwards, armed above and below with numerous small, similar, movable
spines, of which the first four or five are on the carapace ; carapace with antennal
and small pterygostomial spine and a slight dorsal carina on the first half of its length ;
ej-es fairly large and ocellus distinct, but not completely separated from the cornea ;
all the appendages very long and slender; first antenna with rather short stalk,
the first joint longer than the second and third together, the third longer than the
second, both flagella long, the outer broad and flattened at the base, where it bears
a fringe; stylocerite acute; second antenna with the stalk short, and the scale longer
than the antennular stalk, with firm outer edge ending in a spine which arises before
the end of the scale and just projects beyond it; third maxilliped considerably longer
than the antennal scale, with fairly stout meropodite, and the re.st of the limb very weak
and slender ; no epipodite behind the third maxilliped ; first pair of legs considerably
longer than the rostrum ; second pair of legs longer than the antennal scale by the last
half of its wrist ; the latter about 30-jointed, with the last joint equal to two or three
of the preceding joints ; last three legs with the meropodite fairly strong and armed with
spines, and the distal part of the limb very long and weak ; exopodite of uropod longer
than endopodite and armed with a spine on the outside near the fi'ee end ; endopodite
DR WILLEY FROM THE SOUTH SEAS. 413
longer than telson ; telson armed with four pairs of spines above, and with two pairs
at the free end."
2 $ from Blanche Bay, New Britain. 2 j" from the D'Entrecasteau.v Group, British
New Guinea.
28. Pandalus {Pariipandalus) longirostris sp. ii., Figs. IQii — 10/;.
Diagnosis: — " A Piuulalus with the rostrum long, outreaching the antennal scale, bent
upwards and armed above and below with movable spines, those at the base above being
louger and farther apart than those towards the free end, and the first three or four
being placed on the carapace ; carapace with antennal spine, a spine at the pterygo-
stomial angle, and a keel on the anterior half of its dorsal surface ; eyes fairly large,
with ocellus distinct but not completely sundered from the main cornea ; appendages
moderately stout: first antenna with the basal joint of the stalk longer than the
second and third together, third rather longer than second, second covered with hairs,
stylocerite longer than basal joint and ending in a spine, both flagella long, outer
broad and flat at base, in which region it bears a fringe of hairs; second antenna
with short stalk and long scale, whose external tooth barely projects at the end :
third maxilliped slightly longer than the antennal scale ; epipodite wanting from the
last leg only ; first pair of legs very little longer than third maxillipeds ; second pair
of legs equal, with about 25 joints in the wrist ; uropod longer than the telson, which
is narrow, elongated, and armed at the end with one median and four movable lateral
spines, and bears four pairs of spines on the doisal surface."
Length 130 mm.
4 % from New Britain, 2 with eggs.
Genus. Heterocarpus A. M.-Edw., 1881.
29. Heterocarpus ensifer A. M.-Edw., 1881.
Heterocarpus ensifer, A. M.-Edw., Ann. Sci. Nat. (6) xi. 4 p. 8 (1881); Bate,
"Challenger" Macrura, p. 638 pi. cxii. fig. 4 (1888).
In the present specimen the spines on the rostrum vary from 12 to 16 above,
and from 7 to 10 below.
The first leg is simple. In H. gihbusus Bate it is chelate (Caiman, loc. cit.)
3 % and 2 (^ from Blanche Bay, New Britain. 4 young specimens from the
same localitv in 100 fathoms of water.
Family. Hippolytidae.
Genus. Savon Thallw., 1891.
30. Siiron mannoratus (Oliv.). 1811.
See P. Z. S. 1898, p. 1009 (1899).
1 ? from Nivani, Louisiades, British New Guinea.
w. IV. 56
414 0>f the stomatopoda axd ilacruha brought by
Family. Latreutidae.
Genus. Parhippobjte nov.
The absence of a cutting edge (psalistonia) from the mandible of the species on
v,'hich this new genus is founded obliges me to place it in Ortmann's new family,
Latreutidae. [Bronn's " Thierreich," Crust. Ii. p. 1130 (1898).] It is, however, so closely
allied to Merhippolyte Bate that it might almost equally well be placed like the latter
group as a subgenus of Spirontocaris Bate (non Hippolyte Leach, restrict.). In any case,
the difference between the Latreutidae and Hippolytidae will not, I think, be ultimately
found to be of more than subfamily value.
Characters of Parhippolyte n. gen.
1. Rostrum moderate, dentate.
2. Supraorbital spine wanting, antennal and postorbital spines present.
3. Flagella of first antenna long.
4. Mandible without cutting edge, with three-jointed palp.
.5. Third maxilliped with exopodite.
6. Second wrist multiarticulate (about 30 joints).
7. Branchial formula as in Merhippolyte but no pleurobranch on third maxilliped.
8. Sixth abdominal segment with the hinder angle provided with a small spine,
but not articulated.
31. Parhippolyte uveae n. sp., Figs. 11a — llg.
Diagnosis : — " A Parhippolyte with the rostrum, bearing three or four teeth above
and five below; with antennal and postorbital spines present, the pterygostoniial
angle of the carapace rounded and the anterior two-thirds dorsally carinated ; the
antennular stalk half the length of the antennal scale, its first joint almost equal to
the second and third together, the stylocerite equal to the first joint, broad, acute, the
flagella subequal ; the scale of the second antenna long, broad at the base, narrowing
rapidly, with the spine barelj' projecting beyond the free end, flagellum about equal to
the antennular flagella ; third maxilliped outreaching the antennal scale by the last two-
thirds of its last joint, which is obliquely truncated at the end ; first leg not reaching
the end of the antennal scale, hand equal to wrist, fingers shorter than palm, not
dentate, with a small black claw at the tip ; second leg outreaching by the WTist the
antennal scale, \vrist about 30-jointed, first and last joints subequal, about twice the
length of any of the others ; remaining legs long, the last slightly the longest, owing
to increased length of the propodite, meropodite with spines underneath ; endopodite of
iiropod as long as telson, exopodite longer ; telson ending in a median spine and
bearing at the end four lateral spines and on the dorsal surface four jjairs of spines."
Length 110 mm.
Ten specimens, ? all female. Three with eggs. From Uvea, Loyalty Islands.
dr willey from the south seas. 415
Family. Rhtnchocinetidae.
Genus. Rhynchocinetes H. M.-Edw., 1837.
32. Rhynchocinetes typus H. M.-Edw., 1837.
Rhynchocinetes typus, H. Milne-Edwards, Ann. Sci. Nat. 2 vii. p. 165, pi. iv. C. (1837).
The single specimen, which is from Lifu, Loyalty Islands, has on the rostrum only
four spines above and only twelve below.
Family. Alpheidae.
Genus. Alpkeinus nov.
The recent work of Coutiere [Bull. Mus. Paris, il. p. 380 (1896)] on this family
necessitates the establishment of a new genus as well as of a new species for two
specimens of an Alpheid from Lifu.
Characters of Alpheinus n. gen.
1. Eyes completely covered above but not enclosed in front.
2. Rostrum and ocular spines present.
3. Eyestalks short, without spines above. Cornea lateral.
4. Outer flagellum of first antenna slightly bifid at tip.
5. Pleurobranch to each leg. Arthrobranch to third maxilliped. No epipodites.
6. First pair of legs unequal. Left like the large leg of Alpheus, but with movable
finger as in Betaeus. Right small, simple.
7. Angle of sixth abdominal segment not articulated.
33. Alpheinus tridens n. sp., Figs. 12« — 12^.
Diagnosis: — "An Alpheinus with the rostrum of moderate length, shorter than the
first joint of the antennular stalk, triangular with a sharp apex, depressed at base,
compressed at apex, not dentate, with a dorsal keel starting between the eyes ; ocular
spines resembling rostrum but shorter; carapace without other spines than the ocular
and with produced but not acute pterygostomial angle ; first antenna with the stalk
longer than the antennal scale, first joint longer than second and third together, second
longer than third, first two joints projecting on the outside at the distal end and
bearing on the projection a few strong plumose hairs; stylocerite sharp, almost equal
to the first joint ; second antenna with the scale shorter and the stalk longer than
the stalk of the first antenna, scale with strong outer border and freely projecting
spine, basipodite with stout spine on the outer side; third maxilliped very hairy,
reaching the end of the antennal scale ; larger leg of the first pair outreachiug the
antennal scale by the last two-thirds of the palm, hand longer than carapace, fingers
shorter than palm, a spine on the palm at the base of the movable finger, and a tooth
56—2
416 ON THE STOMATOPODA ASD MACRURA BROUGHT BY
on the biting edge of the same finger, \vrist very -short with a spine on the outer
and another on the inner side, meropodite shorter than the palm, with a spine on the
outer side at the distal end ; smaller leg of the first pair longer than the antennal
scale, simple, hairy, with hand long and wrist short, and fingers shorter than the palm ;
second leg outreaching the antennal scale by the last four joints of the wrist, wi'ist
S-jointed, 1=2 + 3 + 4 + 5, 5 = 3 + 4, 2, 3, 4 subequal ; remaining legs rather stout,
propodite longer than carpopodite, shorter than meropodite, carpopodite with one tooth
above at the distal end, dactyle stout, biunguiculate, numerous spines underneath the
propodite ; telson and uropods short and broad ; endopodite and exopodite of uropod
subequal, somewhat longer than telson, exojDod with first joint projecting considerably
outside the second and bearing on the projection a slender spine ; telson with the free
end subtruncate, with a low rounded lobe in the middle, two short spines on each side
and a long fringe, and with two pairs of movable spines on the dorsal surface."
Length 20 mm.
2 specimens from Sandal Bay. Lifu, Loyalty Islands.
Genus. Synalpheus Bate, 1888.
Synalpheus, Bate, Challenger, Macrura, p. 572 (1888) ; Coutiere, Notes, Leyd. Mus.
XIX. p. 206 (1897).
34. Synalpheus biunguiculatus (Stimps.), 1860.
? Alpheus hiunguiculatus, Stimpson, Proc. Ac. N. Sci. Philad. 1860, p. 31.
Alpheus minor, var. hiunguiculatus, de Man, J. Linn. Soc. ZooL, XXII. p. 273 (1888).
Alpheus sp., de Man, Zool. Jahrb. IX. Syst. p. 738, Fig. 62 (1897).
? Alpheus tricuspidatus. Heller, Sitz. Ak. Wiss. Wien, XLiv. p. 267 (1861).
1 $ from the Reef, Ralun, New Britain.
Var. C, nov. One male, and a small specimen with a Bopyrid in the gill chamber,
taken in the mantle cavity of an ascidian at Baravon, New Britain, differ from de Man's
type in having the ocular spines as long as the rostrum and rather broad and triangular.
De Man has named two varieties A and B respectively. I propose to call the present
form var. C.
35. Synalpheus demani nom. nov.
Alpheus triunguiculatu^, de Man, Arch. Xaturg., Liil. 1, p. 508, PI. XXII. Fig. 2
(1887).
According to Coutiere the name triunguiculatus was given by Paulson in 1875 to
a species which must be included m the genus Synalpheus. It is very unlikely that
this species is identical with that to which de Man gave the same name in 1887,
describing it as new. A new name is, therefore, probably wanted, and the most ap-
propriate course is obviously to call the species after its first describer.
2 % from Lifu, Loyalty Islands.
DR WILLEY FROM THE SOUTH SEAS. 417
36. Synalpheus neomeris (de Man), 1897.
Alpheus neomeris, de Man, Zool. Jahrb. IX. Syst., p. 734 (1397).
One specimen, from Sandal Bay, Lifu, Loyalty Islands.
Genus. Alpheus Fabr., 1778.
37. Alpheus ohesomanus Dana, 1852.
Alpheus ohesomanus Dana, U.S. E.^pl. Expeil. Crust, i. p. .574, PI. XXXIV. Fi^^. 7
(1852).
Two specimens from Lifu, Loyalty Islands. One from Blanche Bay, New Britain.
38. Alplteus yracilidigitus Miers, 1884, var.
Alpheus gracilidigitus Miers, "Alert" Report, p. 287 (1884); de Man, Ma.\ Weber's
Zool. Ergebnisse, p. 406, PI. XXV. Fig. 32 (1892).
The specimens differ from the type in that : —
1. The lower border of the merus of the first leg is not serrate.
2. The movable finger of the small chela wants the tooth on the inner side.
One ^ and one % from the Isle of Pines, New Caledonia.
39. Alpheus laevis Randall, 1839.
Fur references see P. Z. S. 1898, p. 1013.
2 $ from Blanche Bay, New Britain. 1 $ from Sandal Bay, Lifn.
40. ? Alpheus diadema Dana, 1852.
? Alplieus diadema Dana, U.S. Expl. Expd. Crust, i. p. 555, PI. XXXV. Fig. 7 a — e
(1852).
Dana describes the first joint of the wrist of the second pair of legs as being
" much longer than the second," but figures it as of almost the same length. In the
present specimen it is very slightly shorter. The hands of the first paii', which were
wanting, from Dana's specimen, are figured from that in the present collection (PI. XXXIX.,
Fig. 17).
1 $ from Sandal Bay, Lifu, Loyalty Islands.
41. Alpheus frontalis Say. 1832.
For references see P. Z. S., 1898, p. 1013.
2 c/" and 1 $ from Lifu, Loyalty Islands.
42. Alpheus aglaopheniae n. sp.. Figs. 13n — 13/!
A single, dismembered specimen of an Alpheus found living among the branches
of a hydroid polyp of the genus Aglaophenia, represents, I think, a new species.
It is diagnosed as follows :
"An Alpheus with the rostrum arising from the anterior border of the carapace,
reaching the end of the first joint of the anteunal stalk, and continued backwards as
a short keel on the carapace ; eye-hoods acute in front ; second and third joints of the
418 ox THE STOMATOPODA AXD MACRURA BROUGHT BY
antennular stalk subequal, first joint somewhat longer than either, stylocerite equal to
the first joint; stalk of second antenna longer than that of the first, scale bearing a
strong spine, equal to the antennular stalk, long fringed; thii-d maxilliped large, covered
with long hairs, projecting beyond the antennular stalk ; larger leg of the first paii-
with the lower border notched but the upper only very faintly so, fingers less than halt
the length of the palm, %\Tist short, somewhat excavated, meropodite broad, with lai-ge
distal spine and distal end excavated, hand hairy; smaller leg of the first pair with hand
elongate, bearing a spine above the movable finger, hairy, fingers equal to the palm,
wrist short, with a spine on the outside, meropodite of the same form as in the larger
hand, but with the spine smaller and the distal end excavate ; second pair of legs with
the first joint of the wrist the longest, 2 and 5 equal, 3 and 4 short, hand about equal
to first wrist-joint; remaining legs without a spine on the meropodite, the propodite
armed with spines, the dactyle biunguiculate, one-third the length of the propodite; the
exopodite of the uropod larger than the endopodite, the latter larger than the telson,
which is hairy above."
Length 11 "5 mm.
One ? from the Engineer Group, British Xew Guinea.
Tribe. LORICATA.
Family. Palinuridae.
Genus. Panidirus White, 1S47.
43. Panulirus demani nom. nov.
PanuUrus polyphagus, Ortmaun, in Semen's " Forschuugsreisen in Austral.," Y. 1,
p. 19 (1894).
Panulirus sp., de Man, Zool. Jahrb. IX. Syst. p. .507 (1898).
There is no evidence for the view that this is a young form of P. poli/phaffus,
and it is therefore well that it should receive at least a provisional name. It is here
proposed to call the species Panulirus demani after the author who first recognised
its distinctness.
One (/, from Blanche Bay, New Britain.
44. Panuli7-us bispinosus sp. u.
A small specimen in the collection seems to deserve a name and a short diagnosis
as a probably new species. It bears a considerable resemblance to P. femoristriga
v. Martens, 1872, of which it may possibly be a young example, but the abdominal
furrows are interrupted in the middle line, and the antennal tergite is quite smooth,
save for two spines towards the anterior edge. P. femoristriga probably also occurs
in the Loyalty Islands, since Dr Willey took, but did not preserve, a large Palinurid
which from his description would seem to belong to that species.
Diagnosis : — " A Panulirus with the stalk of the first antenna somewhat shorter
than that of the second, the first joint longer than the second or thii-d, the third some-
DR WILLEY FROM THE SOUTH SEAS. 419
what lono-er than the second ; the carapace and the stalk of the second antenna covered
with spines of various sizes with their points directed forwards, somewhat sparsely mingled
with hairs ; the antennal segment bearing two spines only and no spinules ; the third
maxilliped with an exopodite bearing a flagellum which reaches half way up the mero-
podite ; the legs hairy, the second pair the longest ; the abdominal furrows interrupted
in the middle line of the body."
Length 25 mm.
One t/ from Sandal Bay, Lifu, Loyalty Islands.
45. Panulirus penicillatus (Oliv.) 1811.
For references see P. Z. S., 1898, p. 1015.
One tf from Natikitiwan, Lifu, Loyalty Islands.
Family. Scyllaeidae.
Genus. ScyUarus Fabr., 1793.
46. ScyUarus sieboldi de Haan, 1850.
Sc>/llarus sieboldi, de Haan, Faun. Japon. Crust., p. 152, PI. XXXVI. Fig. 2 (1850).
1 £/■, 1 $ from Lifu, Loyalty Islands.
Genus. Paribacus Dana, 1852.
47. Paribacus antarcticus (Rumph).
For references see P. Z. S., 1898, p. 1015.
5 (/, 2 $, from Natikitiwan, Lifu, Loyalty Islands.
Tribe. THALASSINIDEA.
Family. Callianassidae.
Genus. CalUanassa Leach, 1814.
48. CalUanassa novae-britanniae sp. n., Figs. 14a — 14cZ.
Diagnosis: — "A CalUanassa with the rostrum short, triangular, not half the length
of the eyestalks; the latter compressed, not quite equal to the first joint of the antennular
stalk, with lateral cornea; carapace with a triangular projection between the eye and
the base of the second antenna, and the pterygostomial region projecting forwards below
the antenna, a well-marked median ridge and cervical furrow, and the hinder border
excavate; first antenna having the second joint of the stalk the longest and the whole
stalk shorter than that of the second antenna; last joint of third ma.xilliped broad and
with a long fringe of hairs, other joints ail fairly broad; first pair of legs unequal,
wrist as broad as hand, fingers shorter than palm, moveable fingers longer than
420 ON THE STOMATOPODA AXD MACRCRA BROUGHT BY
immovable, meropodite armed with spines ; telson short, broader than long, with straight
hinder edge ; uropod longer than telson, with the raised portion of the exopodite not
projecting laterall}- beyond the rest of the structure.
Length 37 mm.
1 (/, from New Britain.
Genus. CaUianidea H. M.-Edw., 1837.
49. CaUianidea bjpa, H. M.-Edw., 1837.
CaUianidea typa, H. M.-Edw.. H. N. Crust, ii. p. 320, PI. XXV. his, Figs. S— 14
(1837).
1 (/ from Sariba, British New Guinea.
Family. Axiidae.
Gexus. Eiconawius Bate, 1888.
50. Eiconaxins taliliensis sp. n., Figs. 15a — 15c.
Diagnosis : — " An Eiconaxiu^ with the rostrum equal to the first joint of the
antennular stalk, ending in two spines, with the sides bent up to form a gutter, and
crowned on each side by spines intermingled with thick tufts of hairs ; on the carapace
this gutter is continued backwards for a short distance, and on each side an interrupted
hairy groove runs back from the base of the rostrum along the sides of the flattened
dorsal area to the cervical furrow, just before meeting which the grooves curve some-
what outwards. The anterior part of the flat area of the carapace is protected at the
side by a raised ridge, which is continuous in front with the side ridges of the rostrum.
Antennal tooth present, and pterygostomial angle produced but rounded ; cervical furrow
deep, and at the side running obliquely into a shallower depression, which continues
it to the anterior edge of the carapace ; outside the cervical groove a small crest of
teeth on each side of the body ; eye-stalks shorter than rostrum, cornea terminal ;
first joint of antennal stalk equal to second and third together, latter subequal ;
stalk of second antenna longer than that of first, scale narrow, strong, with five teeth
beneath, spine on basal joint with four teeth outside and two longer teeth inside ;
third maxilliped longer by its last three joints than the antennal scale, meropodite with
four spines on the inner edge, cai-popodite with five, propodite and last joint nearly equal,
latter elongate-oval ; first pair of legs subequal, left stouter than right but otherwise
similar, ischiopodite with a row of teeth below, meropodite with five or six rather large teeth
above and a crest of small teeth below, immovable finger with one tooth, hand slightly
broader than wrist, which bears a single tooth below ; second pair of legs with a crest
of teeth under the meropodite ; second, third and fourth legs with a ventral process at the
outer end of the ischiopodite ; propodite of legs 3 and 4 with a thick fringe of hairs
below ; all the limbs hairy-tufted ; abdomen longer than cephalothorax ; telson with two
transverse ridges, behind the second ridge two pairs of small tubercles, along the hinder
and lateral borders a row of small, indistinct tubercles, hinder border straight and with
DR WILLEY FROM THE SOUTH SEAS. 421
a fringe of hairs, uropods not longer than telson, with longitudinal ridges on the dorsal
surface of, and toothed on the outer edge of both rami."
Length of male 57 mm., of female 55 mm.
1 £/■, 1 ? from Talili Bay, New Britain.
Tribe. ANOMALA.
SuBTRiBE. GALATHEINEA.
Family. Galatheidae.
Genus. Gulathea Fabr.. 1798.
51. Galathea elegans Adams and White, 1848.
Galathea elegans, Adams and White, Crust., " Samarang," p. 1, PI. XII. Fig. 7 (1848).
1 $, with eggs, from New Britain.
52. Galathea grandirostris Stimps., 1858.
Galathea grandirostris, Stimpson, Proc. Ac. N. Sci. Philad., 1858, p. 1)0; Henderson,
"Challenger" Anomura, p. 119, PI. XII. Fig. 3 (1888).
2 (/ and 2 $, from New Britain.
53. Galathea australiensis, Stimps., 1858.
Galathea australiensis, Stimpson, Proc. Ac. N. Sci. Philad., 1858, p. 89 ; Henderson,
"Challenger" Anomura, p. 118, PI. XII. Fig. 5 (1888).
1 (/ from Lifu, Loyalty Islands.
54. Galathea affinis Ortm., 1892.
Galathea affinis, Ortmann, Zool. Jahrb. VI. Syst., p. 252, PI. XI. Fig. 9 (1892).
2 (/ from Lifu, Loyalty Islands.
55. Galathea spinimanus sp. n., Figs. 16a— IQb.
Diagnosis :— " A Galathea with the rostrum broad, with one small and three large
spines on each side and a terminal spine ; carapace without gastric spines, with six
spines at the side and one at the pterygostomial angle, scored with transverse pilose
ridges, but without demarcation of the gastric region ; third maxilliped with the mero-
podite as long as the ischiopodite, but narrower, and not bearing a spine on the
outside, on the inside of the meropodite two moderately large and two small teeth,
dactyle broad and ending in a tuft of hairs; first pair of legs longer than the thorax,
covered with spines and hairs, fingers about equal to the palm, a small tooth on the
inside of each finger ; second, third and fourth legs covered with spines and haii-s,
dactyle not far short of the propodite in length, with no spines above, but a row of
small spines below."
Length 9'5 mm. df).
1 (/ 2 ? from Lifu, Loyalty Islands.
w. IV. 57
422 ON THE STOMATOPODA AXD MACRURA BROUGHT BY
Genus. Munida Leach, 1820.
56. Munida scabra Hend., 1885.
Munida scabra, Henderson, Ann. Mag. N. H. (5) xvi. p. 409 (1885); "Challenger"
Anomura, p. 134, PI. XV., Fig. 1 (1888).
3 ? from Talili Bay, New Britain.
Var. longipes nov.
A male and two females, taken with the above typical specimens, differ from them
in the greater length and slenderness of the legs of the first pair\ It is proposed to
call this variety longipes.
57. Munida japonica Stimps., 1858.
Munida japonica, Stimpson, Proc. Ac. N. Sci. Philad., 1858, p. 252; Ortmann,
Zool. Jahrb. vi. Syst. 2, p. 254, PI. xi. Fig. 11 (1892).
2 ? from Talili Bay, New Britain.
58. Munida semoni Ortm., 1894.
Munida semoni, Ortmann, Semon's " Forschungsreisen in Austral," v. 1, p. 24, PI. I.
Fig. 4 (1894).
2 J' and 3 ? from Talili Bay, New Britain.
Family. Porcellanidae.
Genus. Petrolisthes Stimps., 1858.
59. Petrolisthes hastatiis Stimpson, 1858.
Petrolisthes hastatus, Stimpson, Proc. Ac. N. Sci. Philad., 1858, p. 241 ; Ortmann,
Zool. Jahrb. vi. Syst. 2, p. 260 (1892).
23 c/" and 14 $ from Sariba, British New Guinea.
60. Petrolisthes lamarchi (Leach), 1820.
See P. Z. S , 1898, p. 464.
Type. 1 c/ and 1 $ from Sariba, British New Guinea.
Var. fimbriatus Borradaile, 1898. 1 ^ and 1 ? from Sandal Bay, Lifu, Loyalty
Islands ; 1 ? from Sariba, British New Guinea.
61. Petrolisthes bispinosus sp. n.
Diagnosis: — "A Petrolisthes with the front indistinctly trilobed, the middle lobe
prominent, each lobe concave above ; carapace covered ^vith straight, continuous, pilose
ridges and bearing on each side two epibranchial spines, but without spines on its
hinder edge ; chelipeds marked out into scales by pilose ridges, their meropodite vnth a
blunt lobe on the inner edge, the wrist with the inner edge 5-lobed, the two proximal
• Taking in each case the average length of the carapace in the specimens before us as unity, the length of
the first pair of legs is in -V. scabra 3--5 and in var. longipes 5"2. The specimens of longipes are slightly smaller
than those of the type.
DR WILLEY FROM THE SOUTH SEAS. 423
lobes each ending in a minute spine, the rest finely serrate, the outer edge with a
crest of sharp, curved teeth, the hand broad, with serrated edges, the fingers slightly
hooked at the tip, equal, with serrated edges ; the second to fourth pairs of legs with
spines on the ujjper edge of the meropodite, dactyies with several small spines under-
neath ; none of the legs hairy, save for a ver}' few scattered hairs."
Length of carapace 4 mm.
This species belongs to the galanthinus-group of Ortmauu [Zool. Jahrb. X. Syst.,
p. 276 (1897)].
1 (/ from Sandal Bay, Lifu, Loyalty Islands.
Genus. Pachycheles Stimps., 1858.
62. Pachycheles scidptus (H. M.-Edw.), 1837.
Porcellana sculpta, H. M.-Edwards, H. N. Crust, ii. p. 2.53 (1837); Dana, U.S.
Espl. Expd. Crust, i. p. 412, PI. XXVI. Fig. 2 (1852); de Man. J. Linn. Soc. Zool.
XXII. p. 218 (1888).
Porcellana pisum, H. M.-Edwards, H. N. Crust, ii. p. 254 (1837); Heller, "Novara"
Crust, p. 73 (1868).
Porcellana pulchella, Haswell, Proc. Liun. Soc. N.S.W. XI. p. 758 (1882); Cat.
Austral. Crust, p. 148 (1882).
Porcellana (Pisosoma) sculpta, de Man, Arch. jSaturg. LIII. p. 413 (1888).
Pachycheles pidchellus, Miers, "Alert" Report, p. 273, PL XXX. Fig. A (1884);
Henderson, "Challenger" Anomura, p. 114 (1888); Ortmann, Semon's " Forschungsreisen
in Austral." v. 1, p. 30 (1894).
Pisosoma sculptum, Ortmann, Zool. Jahrb. VI. S3'st. p. 265 (1892) ; de Man, Zool.
Jahrb. IX. Syst. p. 878 (1896).
Pisosoma jnsum, de Man, Zool. Jahrb. IX. S3-st. p. 380 (1896).
Pachycheles sculptus, Ortmann, Semon's "Forschungsreisen in Austral." v. 1, p. 29
(1894).
The occurrence in Dr Willey's collection of a form intermediate between the Por-
cellana sculpta and P. pisum of Milne-Edwards, leads to the conclusion that these
latter are not specifically distinct, and must rank Axith the new form as varieties of
one species. The following table sets forth the distinguishing characteristics of these
varieties.
1. Chelipeds subequal, similar, tuberculated.
A. var. sculjjtus H. M.-Edw., 1837.
r. Chelipeds unequal, dissimilar, one at least not tuberculated.
2. The left cheliped is the larger. Right cheliped tuberculated.
B. var. tuberculatus nov.
2'. The right cheliped is the larger. Neither cheliped tuberculated.
C. var. jnsum H. M.-Edw., 1837.
57—2
424 OX THE STOilATOPODA AND MACRURA BROUGHT BY
The collection contains the follow-ing specimens : —
var. sculptus. 1 ? from Lifu, Loyalty Islands.
var. tuberculatus. 2 c/' from Lifu, Loyalty Islands.
63. Pachycheles lifuensis sp. n.
Diagnosis : — " A Pachycheles with the front almost straight, slightly convex in the
middle, depressed; carapace granular at the sides and with linear ridges on the branchio-
stegites; chelipeds unequal, the left the larger, the wrist and hand uniformly pubescent
and granular, the wrist with three rather blunt lobes on the inner edge, but without
teeth overhanging its articulation with the hand ; second to fourth pairs of legs not
so stout as in barbatus, the last three joints pubescent, the propodite armed with
spines above, the dactyle with spines below."
Length of carapace 3"5 mm., breadth 4 mm.
This species is closely allied to P. barbatus, but is, I think, distinct.
1 ^ and 1 ? from Lifu, Loyalty Islands.
SuBTEiBE. PAGURINEA.
Fajviilt. Pagueidae.
Genus. Pagurus Fabr., 1793.
64.. Pagurus deformis, H. Il.-Edw., 1836.
For references, etc., see P. Z. S., 1898, p. 460.
The male specimen of this species shows, as usual, the genital openings of the
female.
From Lifu, Loyalty Islands. 1 ? in a Doliuin shell, 1 $ in a Turbo shell bearing
sea anemones, 1 </.
From the Conflict Group, British New Guinea. 1 %. berried, in a Turbo shell,
bearing a sea anemone.
From New Britain ; 2 $ in shells of Doliuin and Natica.
65. Pagurus gemmatus H. 3I.-Edw., 1846.
Pagurus gemmatus, H. M. -Edwards, Ann. Sci. Nat. (3), x. p. 60 (1846).
The male of this species does not show the female openings found in the male
of the allied P. deformis.
1 ^ from New Britain, 1 ^ from Sandal Bay, Lifu, Loyalty Islands: the latter
in a Dolium shell bearing sea-anemones.
66. Pagurus asper de Haau, 1849.
Pagurus asper, de Haan, Faun. Japon. Crust., p. 208, PI. XLIX. Fig. 4 (1849);
Ortmann, Semon's " Forschungsreisen in Austral." v. 1, p. 31 (1894).
1 (/ and 1 ? from Lifu, Loyalty Islands in Turbo shells.
Y Zoological Results
r o P-c dp:
BORRADAILE, CRUSTACEA - MACRURA.
DR WILLEY FROM THE SOUTH SEAS. 425
67. Parjurus setifer H. M.-Edw., 1836.
For references see P. Z. S., 189.S, p. 460.
2 (/ from Lifu, Loyalty Islands. 2 ? from the Conflict Group, British New Guinea.
68. Pagurus euopsis Dana, 1852.
For references see P. Z. S., 1898, p. 461.
1 t/ from the Conflict Group, British New Guinea.
69. Pagurits punctulatus Olivier, ISll.
For references see P. Z. S., 1898, p. 461.
2 (f and 1 ? from Lifu, Loyalty Lslands.
70. Pagurus strigatus (Herbst), 1796.
Cancer strigatus, Herbst, Naturg. Krabben, ii. 4, p. 2.5, PI. LXL Fig. 3 (1796).
Pagurus strigatus, Hilgendorf, Monatsbor. Ak. Wiss., Berlin, 1878, p. 820, PI. II.
Fig. 8.
1 ^ from Sandal Bay, Lifu, LoA'alty Islands.
Family. Coenobitidae.
Genus. Coemhita Latr., 1826.
70. Goenobita compressus H. M.-Edw., 1837.
Coenohita compressa, H. M.-Edw., H. N. Crust. II. p, 241 (1837).
Coenobita compressus, Ortmann, Zool. Jahrb. vi. Syst., p. 318 (1892).
From Lifu, Loyalty Islands, 1 ^ and 2 $, the latter in shells of Nannia and
Papuina. From New Britain, 1 ^ in a Triton shell. From Sandwich Island, New
Hebrides, one berried $ .
71. Coenobita rugosus H. M.-Edw., 1837.
For references see P. Z. S., 1898, p. 460.
1 J" and 12 $ from the Loyalty Islands.
72. Coenobita perlatus H. M.-Edw., 1837.
For references .see P. Z. S., 1898, p. 459.
6 cT and 5 % from Lifu, Loyalty Islands.
73. Coenobita clypeatus (Herbst), 1796.
For references see P. Z. S., 1898, p. 4.59.
2 J" and 4 $ fi-om Lifu, Loyalty Islands.
74. Coenobita spinosus H. M.-Edw., 1837.
For references see P. Z. S., 1898, p. 4-59.
1 (/ in a nutshell of Calophyllum, and 9 $ from Lifu, Loyalty Islands.
426 ON THE STOMATOPODA AND ^ilACRUKA.
Genus. Birgus Leach, 1815.
75. Birgus latro (Linn.), 1766.
For references see P. Z. S., 1898, p. 458.
2 (/" and 8 $ from Lifu, Loyalty Islands.
SUBTRIBE. HIPPINEA.
Family. Hippidae.
Genus. Remipes Latr., 1806.
76. Remipes testudinarius Latr., 1806.
Remipes testudinarius, Latreille, Gen. Crust. Jus. i. p. 45 (1806); de Man, Zool.
Jahrb. IX. Syst., p. 463 (1896).
1 (^ and 2 $ from Sandal Bay, Lifu, Loyalty Islands.
77. Remipes pacificus Dana, 1852.
For references, etc., see P. Z. S., 1898, p. 467.
1 c/, 14 $, from Sandal Bay, Lifu, Loyalty Islands. 7 small c/ from the Isle of
Pines, New Caledonia.
78. Remipes celaeno de Man, 1896.
Remipes celaeno, de Man, Zool. Jahrb. ix. Syst., p. 483 (1896).
2 ^ and 45 ? from Blanche Bay, New Britain. 1 ? from the Isle of Pines.
79. Remipes ovalis A. M.-Edw.
Remipes ovalis, A. M.-Edw., Millard's " Notes sur Reunion," Ann. F., p. 12, PL XVII.
Fig. 5 (1863); de Man, Zool. Jahrb. ix. Syst., p. 471 (1896).
3 ? from New Britain.
80. Remipes admirabilis Thallw., 1891.
Remipes admirahilis, Thallwitz, Abh. Mus. Dresden, p. 36 (1891) ; de Man, Zool.
Jahrb. IX. Syst., p. 466, Fig. 51 (1896).
5 t/' from Blanche Bay, New Britain.
Family. Albuneidae.
Genus. Albunea Fabr., 1798.
81. Albunea microps Miers, 1877.
Albunea microps, Miers, J. Linn. Soc. Zool. xiv. p. 328, PI. V. Figs. 12, IS (1877).
2 J" from Blanche Bay, New Britain.
WiLLEY. Zoological Results
/OoL
ropc </«/
BORRADAILE,CRUSTACEA-MACRURA
£ ty./son. -.afno^'a^t
427
Fig. 1.
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Irt.
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Fig.
2.
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2 a.
11
2 6.
11
2 c.
11
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2/
Fig.
3.
11
3 a.
11
3 6.
11
3 c.
Fig.
4.
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ia.
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4 6.
Fig.
5.
)I
5 a.
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5 6.
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.5 c.
Fig.
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Ga.
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Fig.
7.
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7 a.
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7 6.
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Fig. 0.
EXPLANATION OF PLATES XXXVL— XXXIX.
(Borradaile, Crustacea).
Periclimenes lifuensis Borradaile.
Side view x 6.
Head and carapace from above. INIore highly magnified,
3rd maxilUped. Magnified. The tip of the exopodite is broken off.
Periclimenes tenuipes Borradaile.
Side view x 4.
Head and carapace from above.
3rd maxilliped. Magnified.
2nd „ „
1st ,. „
2nd maxilla. „
More highly magnified.
More highly magnified.
Periclimenes parviis Borradaile.
Side view x 9.
Head and carapace from above,
3rd maxilliped. Magnified.
Periclimenes parasiticus sp. n.
Side view x 10.
Head and carapace from above.
Anchistus hiunguiculatus sp. n.
Dorsal view x 1^.
3rd maxilliped. Magnified.
2nd maxilliped. „
Pontonia ascidicola sp. n. 9 •
Dorsal view x 5.
Lst maxilliped. More highly magnified.
Palaemonopsis willeyi sp. n.
Side view x 3.
Head and carapace from above.
Tail fin from above (Plate XXXVIL).
IMandible. Magnified. The projection on the fore edge in the figure is not part
of a palp but is the base of the molar process.
First antenna.
Pandalus serr-atifrons sp. n.
Side view x 2.
Head and carapace from above x 2.
3rd maxilliped x 3.
End of first leg. Magnified.
Pandalus tenuipes sp. n.
Side view, nat. size. The drawing does not adequately represent the thread-like
appearance of the carpopodite and propodite of tiie legs, nor the fact that
they are slightly swollen at the outer ends. These limbs are in the above
respects unlike those of P. serrati/rons.
428 ON THE STOMATOPODA AND MACRUBA.
Fig. 10. Pandalus longirostris .sp. n.
„ 10 a. Side view, nat. size.
„ 10 6. 3rd maxilliped x 2.
„ 10 c. 2nd „
„ 10 d 1st ,,
„ 10 e. 2nd maxilla.
„ 10/. 1st maxilla.
,, 10^. Left mandible.
„ 10 A. End of 1st leg. Magnified.
Fig. 11. Parhippolyte uveae sp. n.
„ 11«. Side view, nat. size.
„ 116. Head and carapace from above x \\.
,, lie. 3rd maxilliped >; 3.
„ llrf. 2nd
,, lie. 2nd maxilla.
„ 11/. 1st
,, 11(/. Mandible.
Fig. 12. Alpheinus tridens sp. n.
„ \'2a. Dor.sal view ■. 3.
,, 12 6. 3rd maxilliped. Magnified.
12 c. 2nd
,, lid. 1st ,, „
„ 12 e. 2nd maxilla. Magnified.
„ 12/ 1st
,, 12^. Mandible. ,,
Fig. 13. Alpheus aglaopheniae sp. n.
,, 13 a. Side view x 6.
„ 13 6. Head and carapace from above x 10.
,, 13 c. 3rd maxilliped.
,, 13 d. Larger leg of first pair.
,, 13e. Leg of second pair.
„ 13/ „ third
Fig. 14. CalUanassa novae-hritannias sp. n.
,, \\a. Anterior part of body from above x 3.
,, 14 6. Tail fin from above x 2.
„ 14 c. 3rd maxilliped x 3.
,, 14 d Larger leg of first pair x 2.
Fig. 15. Eiconaxius taliliensis sp. n.
„ 15 a. Side ^•iew x 2.
„ 15 6. Head and thorax from above x 4.
,, 15c. Tail fin from above x 2.
Fig. 1G. Galalhea spinimanus sp. n. Dorsal view of Ijodv x 6.
Fig. 17. Alpheus diadema Dana.
,, 17 a. Larger leg of first pair x 7.
„ 17 6. Smaller , . x 9.
WiLLEY- Zoological Results
XXXVill
BORiv\UAiLi:;,c;ilUS
£. Wffsoft. Cam6fta'0e
'rtUKA.
WiLLEY Zoological Results.
Plate XXXIX.
ro p-c. at/.
BORRADAILE,CRUSTACEA-MACRURA.
C. iV//sof7, Ctim6^/<f^e.
REPORT ON THE SLUGS.
By WALTER E. COLLINGE, F.Z.S.,
Mason University College, Birmingham.
With Plates XL. and XLI.
PAGE
L Introduction 429
n. Species from the Loyalty Islands 431
1. Veronicella willeyi, sp. nov.
2. Aneitea hirudo ?, P. Fi.soh.
III. Species from the New Hebrides 435
1. Veronicella hrunnea, .sp. nov.
2. „ leydigi, Simr.
3. ,, hedleyi, Simr.
IV. Species from New Britain 436
1. Aiieitella herghi, Plate,
var. nov. alhida.
var. nov. fuscopallescens.
I. INTRODUCTION.
The coUectiou of slugs made by Dr Willcy, although not a large cue, is of
exceptional interest in that it includes a series from a region which as yet has onh-
been very imperfectly worked.
The specimens which are here described belong to two families, — Veronicellidae
and Jauellidae — and include six species of which two are new, and two varieties, both
of which are new.
w. IV. 58
430 REPORT OX THE SLUGS.
So far as I am aware there are no previous records for any species of Veronicella
from either the Loyalty Islands or the New Hebrides, and Professor Simroth informs
me that he also knows of no such records. It is interesting to find amongst the
specimens collected in the New Hebrides, the two Australian species V. leydigi, Simr.,
and V. hedleyi, Simr.
In dealing with a family of molluscs like the Veronicellidae, one is very forcibly
impressed with the little value that can be attached to the external form and colour.
In spite of all the arguments in support of describing and identifj'ing such molluscs
from these features, here they are quite secondary, and to attach to them any special
importance would only lead to endless confusion; a reference to the works of Semper
and Simroth will illustrate the great similarity in colour and external markings that
exists, in species which are widely separated both geographically and anatomically.
Thanks to the work of the above-mentioned malacologists, we have anatomical details
for a large number of species in this family, and in separating those here enumerated
I have been guided almost entirely by their structure. It is much to be regretted
that we have as yet no account of the developmental history of some species of
Veronicella, for such a study would, I strongly believe, throw much light upon their
systematic position and their affinities to other families.
At present it is difficult to say what characters may be regarded as affording
the best criteria for specific distinction, the chief points which I have here directed
attention to, where the material has permitted, are those enumerated by Simroth', viz. :
a. The relations of the intestine to the liver.
h. The differences in the distance between the terminal portion of the intestine
and the female genital orifice.
c. The form and structure of the pedal gland.
d. The form of the salivary glands.
e. The terminal ducts of the male generative organs, particularly the recepta-
culum seminis and vas deferens.
/. The thickness and structure of the notum.
Of the second family, the Janellidae, there are two species, one of which has
formed the subject of part of an elaborate memoir by Plate^, the other is an immature
example of Aneitea, probably A. hirudo, P. Fischer'.
I desire to express my best thanks to Professor H. Simroth for kind assistance,
and to Professor L. H. Plate, who very kindly sent me the type specimen and dissected
parts of Janella schaunislcnidi Plate, from the Bremen Museum, for comparison.
1 Zool. Jahrb. {Abth. f. Syst.), 1890, Bd. v, p. 902.
2 Zool. Jahrb. (Abth. f. Anat.), 1898, Bd. ii, pp. 193—280, Taf. 12—17.
3 Journ. de Conchyl., 1868, T. xvi, pp. 145—46, 225—34, pi. xi.
KEPORT OX THE SLUGS. 431
II. SPECIES FROM THE LOYALTY ISLANDS.
1. Veronicella willeyi, sp. nov. (Pis. XL. — XLI. Fig.s. 1 — 14).
Habitat. Lifu, Loyalty Islands. Numerous.
The colour of this species is exceedingly variable, the majority of specimens were
a dirty yellowish-brown dorsally, irregularly blotched with black, which markings become
more closely set laterally, forming a broken line ; there is a well-marked median dorsal
yellowish-browu line, while the extreme anterior and posterior portions of the body are
a dark bluish-black. The two specimens figured (PL XL. Figs. 1 and 3) show the dark
and light coloured forms. The hypnotum and foot-sole are almost white.
Length (in alcohol) 48 mm. ; foot-sole 5 mm. broad ; hypnotum G mm. broad ;
female generative orifice on the right side 1'.5 mm. from the foot-sole, 27"5 mm. from the
right lower tentacle, and 20'5 mm. from the posterior end of the body.
I have pleasure in associating with this interesting species the name of Dr Willey.
ANATOMY.
I. Digestive System.
The mouth is somewhat oval shaped. The buccal cavity calls for no special
mention. The salivary glands are profuse and lie at the anterior part of the pharynx,
and upon the dorsal side of the buccal cavity. The right gland is slightly smaller
than the left one, a feature, I believe, common to all species of Veronicella, in
consequence, as pointed out by Simroth', of the anterior portion of the male organs
restoring symmetry. To the naked eye the separation between the two glands is scarcely
distinguishable ; under the microscope, however, each is seen to consist of a series of
fine dendritic tubes, each of which terminates in one or more small, flat, sac-like
bodies (PI. XL. Figs. 6 and 7). The whole mass is very compact and forms a conspicuous
yellowish-white body above the hinder portion of the buccal cavity and the anterior
portion of the pharynx (PI. XL. Figs. 5 and 6).
The oesophagus is short, giving place to a wide, thin-walled crop (the fore-stomach
of Simroth and others) which becomes constricted just in front of the stomach
(PI. XL. Fig. 4). The disposition of the intestinal loops is very much the same as in
V. hennigi, Simr. (I.e. (1) Taf. XLix. Fig. 12), though in form and structure the digestive
tract is more closely related to V. hedleyi, Simr.
We may conveniently divide the intestinal canal into four loops, the first extending
from the buccal cavity to the stomach, the second from the stomach to the anterior
lobe of the liver, the third lying superficially in the liver, from the anterior lobe, and
making a somewhat [-shaped bend, which, on leaving the liver, becomes loop number
four, this terminating at the cloacal chamber. The third loop lies dorsal to the crop
• T. c, p. 86G.
58—2
432 REPORT OX THE SLUGS.
imbedded in the substance of the liver, and enters as loop number four into the body
wall just behind the opening of the female generative orifice. The anterior portion
of loop four passes backwards above the kidney, posteriorly it is covered by the
body-wall only, and terminates on the ventral side of the cloacal chamber (PI. XL.
Figs. 9 and 11).
The whole of the intestinal tract, it will thus be seen, is practically imbedded in
the liver, so that V. %uilleyi in this particular agrees with that group of Veronicella
which would also contain V. hennigi, Simr., and V. hedleyi, Simr.
I was not able to detect any ring-like swelling between the oesophagus and crop,
but between the terminal portion of the crop and the commencement of the stomach
there is a thick muscular ring-like constriction, immediately behind which the hepatic
ducts open.
II. The Pedal Gland.
Although approaching somewhat the condition which obtains in V. leydigi, Simr.,
there are a few well-marked features in the gland of V. willeyi in which it
differs from the former.
The gland commences as a slit-like opening, e.g. O, immediately above the foot-
sole, the anterior edges of the latter partly hiding the opening of the gland. Lip-like
protuberances form the boundaries of the commencement of the ca^'ity. The gland
lies free in the body-cavity upon the dorsal side of the muscles of the foot-sole.
The anterior portion is slightly wider than the rest of the gland, agreeing in this
particular in all the specimens dissected. In general shape and size considerable
differences were noted (PI. XL. Fig. 8 a — c) ; in all the specimens, however, some
portion, either to the right or left, was turned forwards (PI. XL. Fig. 8 a, b, c).
In transverse section the lumen of the gland is almost circular, the glandular
wall being about twice the width of the lumen. The epithelial lining of the gland
is continuous around the cavity.
III. The Kidney, Lung and Pallial Organs.
In V. willeyi I have been able to trace the ureter, and have figured in some
detail the relations of kidney and ureter with the lung and the hinder part of the
intestine (PI. XL. Figs. 9 to 11).
The general structure of these organs calls for no special mention excepting that
here the trabecular tissue is very dense and much folded. In general outline the
kidney is not unlike that in V. leydigi, Simr., only differs in extending for some
short distance below the hinder part of the intestine; posteriorly it narrows and opens
into the rectum by a short but wide ureter which is plainly visible under a low-
power dissecting microscope. Towards the outer wall of the pericardium and on the
dorsal side is a minute but very distinct slit-like opening, which seems to be connected
with the anterior and inner portion of the kidney. Possibly this is the reno-pericardial
opening, but as I was unable to verify this by the transverse sections, it must be
regarded only as a supposition that this is the true reno-pericardial opening. On the
REPORT ON THE SLUGS. 433
outer side of the kidney is the lung, which also proceeds very far backwards, extending
to a point just beyond the terminal portion of the kidney (PI. XL. Fig. 9). The
terminal portion of the respiratory duct opens separately from that of the combined
rectum and ureter, and slightly above it, so that there is a distinct cloacal chamber
into which these two ducts open (PI. XL. Fig. 11). The external opening of this
chamber lies on the right side of the hypnotum (PL XL. Fig. 2).
IV. TJie Generative System.
Although conforming in general to the type of reproductive organs common to
Veronicella, there are certain well-marked features which have specific value.
The hermaphrodite gland is pyriform in shape, and has a closely convoluted duct
which is enclosed in a fine membranous sheath (PI. XL. Fig. 12, h. d.) at its base,
and lying upon, and in life completely hidden by, the albumen gland, is a small
diverticulum, the vesicula seminalis (PI. XL. Fig. 12, v. s.). The albumen gland is
a loose glandular organ not unlike that in V. leydigi, Simr. The oviduct is a long
wide tube, twisted many times upon itself. The middle portion is the widest, while
its commencement is the narrowest ; it also becomes narrow again before opening
externally. The vas deferens is a narrow tube, and bears at its lower end a large
spermatocyst, which has a duct of some considerable length (PI. XL. Fig. 12, sp.).
At the opposite side of the vas deferens, at the point where the duct of the
spermatocyst enters, a duct is given off to the receptaculum seminis. The form and
position of the spermatocyst, together with its duct, is very characteristic of this
species and separates it from any allies. The receptac^dum seminis is a large ovoid
body, with a long tivisted duct (PL XL. Fig. 12, rs., rd.).
In the male organs I follow Simroth in terming the penis-gland of Semper, the
dart-gland or dart-sac.
The penis is enclosed in a thick muscular sac, and in general outline and structure
(PL XL. Fig. 13, p.) agrees very closely with that figured by Simroth for V. lei/digi\
being long and conical in shape and having an oblique slit at the top; there is,
however, in V. willeyi, quite a large vestibule into which the penis opens, slightly
above the opening of the dart-sac. Attached to the penis, just below the point of
origin of the vas deferens, is a strong retractor muscle, and a smaller one just below
the vestibule (PI. XL. Fig. 13, rm., rm.'). The dart-sac is rather longer and \\ider
than the penis (PL XL. Fig. 13, ds.). At the base of the sac there is a small
conical papilla, the dart (PL XL. Fig. 13, d.), and connected with this, but outside
the actual dart-sac, are a series of long thread-like accessory glands (PL XL. Fig.
13, ac. gl.) ; these all open at the base of the papilla or dart, the opening at the
apex of the dart forming a common orifice. The number of these accessory glands was
not constant, the average is 15, but in one case there were only 12, while in another
there were as many as 20.
Whether or not this dart-gland is homologous with the dart-sac in other
Pulmonates, e.g. Parmarion, Bamayantia, or with the dart-sacs in certain species of
Helix and Zonites, which contains an imperforate calcareous dart, I at present hesitate
' Cp. T. c. Tiif. L, Fig. 4.
434 REPORT ON THE SLUGS.
to say. Simroth certainly thinks they are, for he writes {I.e., p. 879), "Die Summe
dieser Dateu macht es wohl mehr als wahrscheinlich, dass auch bei den Vaginuhden
das Reizorgan als Liebespfeil zu deuten sei."
V. The Integument.
The body-wall of the dorsum or notiim is somewhat thin; in transverse section
it is seen (PI. XLI. Fig. 14) to consist of an outer epidermal layer, immediately
beneath which is a dense layer of blackish pigment. This layer also borders the
sides of the mucous glands; of these latter I have obtained very good sections by
freezing, staining with an aqueous solution of magenta, and mounting in a glycerine
fluid. The lumen of the gland is narrow, becoming in must cases slightly larger at
the base; around this canal are a series of bladder-like cavities, which probably serve
as reservoirs for the mucous.
Excepting that the notum in V. ivilleyi is thinner than that described by Simroth
in V. leydigi, my observations agree practically in all particulars with his. In the
mucosa, the concretions which Simroth thinks may be uric acid, were very plentiful
and in a few cases very large.
2. Aneitea hirudoh P- Fisch. (PI. XLI. Figs. 15—17.)
Hab. Lifu. One specimen.
After a careful comparison of this small species with all the described forms,
I am placing it with a query under A. hirudo, P. Fisch. From the imperfect
development of the generative organs I conclude it is a young example. It agrees
with Fischer's figure' of A. hirudo in the form of the penis (PL XLI. Fig. 16, p.).
The whole of the body is a dirty greyish-brown colour, vnth a few blotchy black
markings on the lateral portions of the body. There is an ill-defined median-dorsal
groove with oblique lateral grooves directed posteriorly and ventrally. The margin of
the dorsum (perinotum) is wavy. The foot-sole is marked by a series of closely set
transverse markings. In transverse section the body appears almost triangular (PI. XLI.
Fig. 17).
The dimensions are as follows : —
Length over back, from head to tip of the tail 22'.5 mm.
Length of foot-sole 19'5 mm.
Width over back 10 mm.
Breadth of foot-sole 6'5 mm.
Distance of anus from right tentacle 6'.5 mm.
„ „ „ respiratory orifice 1 mm.
Length of mantle 7 mm.
1 Journ. de Conchyl., 1868, T. xvi, pi. xi.
REPORT ON THE SLUGS. 435
III. SPECIES FROM THE NEW HEBRIDES.
1. V. briinnea, sp. nov. (PL XLI. Figs. 18—23).
Hab. Esafate, one specimen.
This is a very interesting form, and I regret, owing to lack of material, not to
be able to give a fuller account of its anatomy.
Externally it is a deep brown with small black blotches sparingly distributed over
the dorsum (notum), the perinotum is very prominent and much darker in colour.
I know of no other species of Veronicella in which the perinotum is so prominent
and so well defined, standing out quite distinct from either the notum or hypuotum.
My knowledge of the genus is not a wide one, so possibly this is a feature met Avith
in other species, still I find no mention of it in any of the species described by
Semper or Simroth. The hypnotum is rather lighter in colour and free of any markings ;
foot-sole yellowish-brown, and marked by a series of transverse divisions.
Length (in alcohol) 18.5 mm.; foot-sole 3 mm. broad; hypnotum 4o mm. broad;
female generative orifice on the right side 1"5 mm. from the foot-sole, 10'.5 mm. from
the right lower tentacle and 7 mm. from the posterior end of the body.
The only parts I have examined anatomically are the digestive system and pedal
gland. The former, excepting in the position of the looiDS of the intestine and stomach,
calls for no special mention. All the loops are imbedded in the lobes of the liver,
those visible on the dorsal surface being the anterior portion of loop 2, and nearly
the whole of loop 3 (PI. XLI. Fig. 19). At the posterior portion of the stomach, and
on the ventral side, is a small bean-shaped glandular body, connected by a series of
fine ducts (PL XLI. Figs. 20 — 22). I have not previously met with any similar
body in any other species of Veronicella. Possibly it functions as a digestive gland.
The pedal gland commences as a wide thin non-glandular (?) sac, lying free in the
body cavity, giving place in the posterior half to a narrower glandular part. At its
extreme end it makes a bend to the right side (PL XLI. Fig. 23).
2. V. leydigi, Simr.
Hab. Esafate, two specimens.
I have nothing to add to the admirable account given by Simroth' of this species.
One specimen is immature.
3. V. hedleyi, Simr.
Hab. Esafate, one specimen.
This specimen agrees in nearly all particulars with the figure made by Hedley
and given by Simroth ^
1 T. c, pp. 865—899. - T. c, Taf. slix, fig. 7.
436
REPORT OX THE SLITGS.
IV. SPECIES FROM NEW BRITAIN.
1, Aneitella berghi, (PI. XLI. Figs. 24—27).
Hab. Karavia, Gazelle Peninsula. Numerous.
This species was first described by Plate' in 1898, who gives numerous figures
of the internal structure, but the external features are scarcely done justice to. Seeing
how very imperfectly figured most of the Janellidae are, a fact I have previously
drawn attention to", I have given two figures of the external form (PI. XLI. Figs.
24—2.5).
The specimens investigated by Plate were from Stephen's Island, New Zealand.
All the specimens I have dissected show a little variation in the form of the
generative organs from those figured by Plate (I. c, Taf. 16, Fig. 55). In one specimen
the penis exhibited a well-marked fold, as figured (PI. XLI. Fig. 27).
The dimensions of the largest specimen are : —
Length over back, from head to extreme posterior 48'5 mm.
Length of foot-sole 45 mm.
Width over back 24 mm.
Breadth of foot-sole 7 mm.
Distance of anus from right tentacle 9"5 mm.
„ „ „ respiratory orifice 2 mm.
Length of mantle 9 mm.
With the typical examples were two well-marked colour variations, which are
described below.
Var. nov. albida.
Hab. Karavia, New Britain, two specimens.
W^hole of body a pure white. Length (in alcohol) 46 mm.
Var. nov. fuscopallescens.
Hab. Karavia, New Britain.
Whole of animal a pale brownish-yellow, with little, if any, black mottling.
Length (in alcohol) 38 mm.
1 T. c, p. 197.
Proc. Zool. Soc, 1894, p. 530.
WujL.by Zoological Results.
Plate Y'-
1\
/m-^\
l^k
*
^.
%,^yj/
1^ ..,
<^T ' \ !l I I ./
WJi.C del.ad.nat
Weot, Newnum lidi.
COL.L.1NOE. SLUGS.
REPORT ON THE SLUGS. 437
EXPLANATION OF PLATES XL. AND XLI.
(Collinge, ShiffS.)
Yeronicella willeyi, sp. nov.
Fig. \. View from dorsal side. Nat. size.
Fig. 2. View from ventral side. Nat. size. 9 female generative orifice, cl. opening of
cloacal chamber.
Fig. .3. Light coloured form, view from dorsal side. Nat. size.
Fig. 4. Intestine and liver, x 2 '5. ce. oesophagus, cr. crop. st. stomach, int^. — int*. the
four loops of the intestine. I. liver. The dotted portion indicates the terminal portion of
the intestine imbedded in the body wall.
Fig. 5. Salivary glands as seen from the dorsal surface.
Fig. 6. Portion of the same, slightly magnified, showing the salivary ducts, s. d. and
their position in relation to the pharynx, j)]i. and tlie buccal cavity, h. c.
Fic;. 7. Terminal portion of the same, showing sac-like bodies.
Fig. 8. Pedal gland showing variations met with, a, h, and c. x 2.
Fig. 9. Mantle organs seen from the venti-al side. atr. atrium, p. pericardium, v. ven-
tricle, h. kidney. I. lung. cl. cli. cloacal chamber, res. d. respiratory duct. iir. ureter.
Fig. 10. Terminal portions of the intestine, kidney and lung. Lettering as in Fig. 9.
Fig. 11. Terminal portions of the intestine and respiratory duct, showing their openings
into the cloacal chamber cl. ch. a. anus. r. o. respiratory orifice, int''. posterior portion of
the intestine.
Fig. 12. Generative organs, alb. gl. albumen gland, h. d. hermaphrodite duct. h. gl.
hermaphrodite gland, ov. oviduct, r. d. receptacular duct. r. s. receptaculum seminis. sp.
spermatocyst. v, d. vas deferens, v. s. vesicula seminalis.
Fig. 13. Terminal ducts of the male generative organs, ac. gl. accessory glands, d. dart.
d. s. dart-sac. p. penis, r. m. and r. m. retractor muscles of the penis, v. vestibule, v. d.
vas deferens.
Fig. 14. Transverse section through the dorsum, ep. epidermis, gl. glands, mu. mucous
layer containing uric acid (?) concretions u. c. m. f. muscle fibres.
Aneitea hiriido, P. Fisch.
Fig. 1.5. Lateral view of the animal from the right side, x 2.
Fig. 16. Vagina vg. and penis p. of the same.
Fig. 17. Diagrammatic transverse section through the body, x 1.
Veronicella brunnea, sp. nov.
Fig. 18. View from the dorsal side, x 2.
Fig. 19. Intestine and liver, x 4. Lettering as above.
w. IV. 59
438
REPORT ON THE SLUGS.
Figs. 20, 21. Dorsal and ventral \-iew of the stomach, the latter showing the position
of the small bean-shaped gland.
Fig. 22. Bean-shaped gland detached.
Fig. 23. Pedal gland.
Aneitella berghi, Plate.
Fig. 24. View from the dorsal side, x 2.
Fig. 25. The same, from the ventral side, x 2.
Fig. 26. Generative organs.
alh. gl. albumen gland.
f. ov. free oviduct.
gl. gland (?)
h. d. hermaphrodite duct.
h. gl. hermaphrodite gland.
p. penis.
pr. prostate.
Fig. 27. Variation in the form of the penis.
r. m. retractor muscle.
r. s. receptaculum seminis.
v. vestibule.
V. d. vas deferens.
vg. vagina.
V. s. vesicula seminalis.
Wiijley Zoological Res-ults.
Plate XL I.
.. N^
IG
n
18 -2
H
■SOtfi.
wit?
-^..
19
24 «z
V^
^ (
23
>mJ:
27
■WEC.del.aviiat.
.^J.Parlrid^e clelri^.14-.
30
. ^
/ I) '
\
26 ft jl
•ts'-U
21
25 « 2 "V=*
Weflt.'Newinan litla
COlililNGE. SLUGS.
REPORT ON THE POLYZOA COLLECTED BY DR WILLEY FROM
THE LOYALTY LSLES, NEW GUINEA AND NEW BRITAIN.
Bv E. G. PHILIPPS,
Newnham College, Cambridge.
With Plates XLII. and XLIII.
The Polyzoa collected by Dr Willey were obtained mainly from Lifu, in the
Loyalty Isles.
The collection contains 63 species of which 9 are new. They are divided among
the three sub-orders as follows :
Cheilostomata 46 species.
Cyclostomata 16 species.
Ctenostomata 1 species.
The specimens were compared with those in the British Museum and in the Cam-
bridge University Museum of Zoology, to which Dr Willey has presented his collection.
I wish to express my thanks to Mr Kirkpatrick for facilities afforded me at the
British Museum and especially to Dr Harmer for the interest he has taken in my
work and the very kind help he has given me.
The nomenclature adopted in this paper is that used in Miss Jelly's " Synonymic
Catalogue of Marine Bryozoa," in which full references are given to the literature on
the group.
Cheilostomata.
1. Catenaria otophora, Kirkp. D'Entrecasteaux Group, Briti.sh New Guinea.
2. Caberea lata, Busk. Sandal Bay, Lifu, 17 fathoms.
3. Cauda retiformis, Pourt. Sandal Bay, Lifu, 80 to 40 fathoms.
4. Scrupocellaria scrupea. Busk. Lifu.
5. Scrupocellaria macaudrei. Busk. Sandal Bay, Lifu.
6. Scrupocellaria annectens MacGill. Karakoai. New Britain, 2 to 3 fathoms.
59—2
440 REPORT OX THE POLYZOA COLLECTED BY DR WILLEY
7. Didymia triserialis, n. sp. Beach of the He du Phare, Noumea, New Caledonia.
8. Bugula dentata, Lamx. Lifu.
9. Bugiila a^-icularia, Linn. Sandal Bay, Lifu.
10. Bugula neritina, Linn. Sandal Bay, Lifu.
11. Tubucellaria cereoides, Ell. and Sol. Lifu.
12. Cribrilina radiata var. ^, Hincks. Sandal Bay, Lifu, 17 fathoms.
13. Membranipora radicifera, Hincks. var. intermedia, Kirkp. Sandal Bay, Lifu,
30 to 40 fathoms.
14. Membranipora lacroixii, Aud. Blanche Bay, New Britain.
15. Membranipora irregularis, d'Orb. Blanche River, New Britain.
16. Membranipora coronata, Hincks. Lifu.
17. Membranipora punctigera. Hincks. Sandal Bay, Lifu.
18. Micropora sp. Lifu.
19. Monoporella pol3Tiiorpha, n. sp. Lifu.
20. Monoporella spinifera, n. sp. Lifu.
21. Schizoporella biaperta, Mich. Lifu.
22. Schizoporella torquata, Q. and G. Beach of the He du Phare, Noumea, New
Caledonia.
23. Schizoporella triangula, Hincks, Blanche River, New Britain, 40 fathoms.
24. Schizoporella striatula, Smitt, Lifu.
25. Schizoporella sanguinea, Norman, Lifu.
26. Schizoporella nivea. Busk. Beach of He du Phare, Noumea, New Caledonia.
27. Schizoporella depressa, n. sp. Lifu.
28. Hippothoa divaricata, Lamx. Lifu.
29. Thalamoporella rozieri, form indica, Aud. Deboyne Lagoon, Louisiades.
30. Microporella ciliata var. personata, Busk. Lifu.
31. Adeonellopsis violacea var. plagiopora, Hincks. Sandal Bay, Lifu.
32. Lepralia feegeensis. Busk. Lifu.
33. Lepralia poissonii, Aud. Lifu, 35 fathoms.
34. Lepralia tuberculata, n. sp. Lifu, 35 fathoms.
35. Lepralia calyciformis, n. sp. Lifu.
36. Smittia marmorea, Hincks. Sandal Bay, Lifu, 35 fathoms.
37. Mucronella articulata, n. sp. Sandal Bay, Lifu, 30 to 40 fathoms.
38. Rhj-nchozoon bispinosum, Johnst. Lifu.
39. Rhynchozoon crenulatum. Waters. Lifu.
40. Escharoides spinigera, n. sp. Lifu, 35 fathoms.
41. Cellepora speciosa, MacGill. Sandal Bay, Lifu.
42. Cellepora mamillata, Busk. Sandal Bay, Lifu.
43. Cellepora simplex, MacGill. Sandal Bay, Lifu.
44. Cellepora longirostris, MacGill. Lifu.
45. Retepora phoenicea, Busk. Sandal Bay, Lifu.
46. Retepora denticulata, Busk, Lifu.
FROM THE LOYALTY ISLES, NE^V C4UINEA AND NEW BRITAIN. 441
Cyclostomata.
47. Crisia aculeata, Hassall, Lifu.
48. Crisia denticulata, Lamk. Lifu.
49. Tubulipora, sp. Lifu.
50. Idmonea interjuncta, MacGill. Lifu.
51. Idmonea radians, Lamk. Lifu.
52. Idmonea australis, MacGill. Lifu.
53. Entalopora delicatula, Busk. Sandal Bay, Lifu, 17 fiithoms.
54. Hornera spinigera, Kirkp. Lifu.
55. Lichenopora verrucaria. Fabric. Lifu.
56. Lichenopora holdsworthii, Busk. Lifu.
57. Lichenopora hispida, Fleming. Lifu, 17 fathoms.
58. Lichenopora ciliata. Busk. Lifu, 17 fathoms.
59. Lichenopora californica, D'Orb. Sandal Bay, Lifu.
60. Lichenopora truncata, n. sp. Lifu.
61. Radiopora cristata, Busk. Lifu.
62. Fasciculipora carinata, Ortmann. He du Phare, New Caledonia.
Gtenostomata.
63. Zoobotryon pellucidum, Ehrenberg. Isle of Pines.
1. Catenaria otophora, Kirkpatrick.
Kirkpatrick, Ann. Mag. Nat. Hist. (6) V. 1890, p. 17.
A few colonies of this delicate species occur, growing on the Alcyonarian
Clavularia viridis.
2. Caberea lata. Busk. Busk, Brit. Mus. Cat. i. 39. Fairly common in Lifu.
3. Canda retiformis, Pourtales. (PL XLIL, fig. 1.)
Cauda retiformis, Pourt. Bull. Mus. Zool., Camb., U. S., 1867, Vol. I., p. 110.
Caberea retiformis, Smitt. " Floridan Bryozoa" in Kongl. Svenska Vetenskaps-Akad.
Handl. 1872—1873, p. 16.
Compare Canda arachnoides, Brit. Mus. Cat. i. 26 and Busk, Challenger Report, XXX.
25.
The specimens, which are common in Lifu, seem to be intermediate between
Canda retiformis, Pourtalfes and Canda arachnoides, Lamouroux. The foiiiix is \w\\
developed, its lamina being as long as the aperture, rounded at the base, produced
and pointed above. It differs in this respect from the specimen of Canda, retiformis
(from Victoria Bank, Brazil) in the British Museum, in which the lamina is small
and equally developed on both sides of the stalk. Smitt however (Flor. Bryozoa, I.
PL v., fig. 44) figures the fornix in this species as varying considerably' in form.
On each side of the upper margin of the aperture there is a spine, the two
being equally developed.
442 REPORT ON THE POLYZOA COLLECTED BY DR WILLEY
The specimen from Brazil, moreover, has no avicularia and this appears to be
characteristic of Pourtales' form. The Lifu specimen has prominent avicularia along
the median line of the branch. The avicularian cell is very large with a serrated
edge : the mandible is triangular and acute.
The ovicells are tj'pical, having the form characteristic of both species. Each
ovicell is crowned by a sessile avicularium and has a large membranous area in front.
The collection contains some other specimens closely resembling the one already
described, but in these the fornix has a small lamina equally developed on both sides
of the stalk, while the mandible of the avicularium is longer.
It will thus be seen that the Lifu specimens resemble Canda arachnoides and
differ from Cauda retifonnis in possessing avicularia and two spines on the upper
margin (in Smitt's figures, most of the zooecia bear only one spine) whereas it differs
from Canda arachnoides in the possession of a fornix and in the greater size of the
avicularium.
4. Scrupocellaria scrupea, Busk. Busk, Brit. Mus. Cat., i. 24.
Only a few colonies occur.
5. Scrupocellaria macandrei, Busk. Busk, Brit. Mus. Cat. i. 24.
Only one small piece occurs.
6. Scrupocellaria annectens. MacGillivray. MacGill. Trans. Roy. Soc. Yict., 1886,
p. 184.
There are two small, somewhat worn fragments which appear to belong to
MacGillivray's species. They agree with it in the general character of the zoarium,
the form of the zooecium and of its orifice, the character of the anterior avicularia,
and in the absence of the fornix in sterile zooecia. Here as in Scrupocellaria
annectens, there is a single vibraculum in the angle of bifurcation of the branch, and
the anterior avicularium at the base of the zooecium lying at the point of bifurcation
is larger than the others. The specimens differ from MacGillivray's species in possessing
three spines at the external angle of all the cells in addition to the spine placed at
the internal upper angle. The vibracula here seem to lie somewhat nearer the
middle line, and the apex of the vibracular cell from which the seta projects, stands
out as a prominent conical process. This feature is not shown in MacGillivray's
figure. No lateral avicularia are seen on the specimens.
7. Didymia triserialis, n. sj). PL XLII., figs. 2 and 2 a.
Cells joined side to side, varying from four to twenty-eight in each internode,
the ordinary zooecia being arranged biserially. At a bifurcation each cell of a primary
pair gives off a secondary pair at its summit ; ovicell smooth, imperforate, divided
by curved lines into three areas, occupying about half the length of the cell, the
orifice being a little above the centre ; ooecial cells placed one above the other
along the middle line of an internode interposed between the two lines of ordinary
zooecia.
FROM THE LOYALTY ISLES, NEW GUINEA AND NEW BRITAIN. 443
Measurements : zooecia from '69 mm. to '89 mm. in length x '26 mm. in width.
ooecia "24 mm. x "24 mm.
The position and structure of the ovicell are the characteristic features which
differentiate this species from Didymia drnplex, Busk, (Brit. Mus. Cat. I. p. 35). The
ooecial cell does not occur at the bifui-cation of a branch as in Busk's species, but
in the middle line of an internode interposed between two ordinary zooecia, so that
here the branch has three cells at the same level. There are several ovicells in an
internode placed one above the other, and occasionally the fertile cells seem to be
succeeded by an additional line of sterile zooecia so that the branch becomes triserial.
This apparently occurs at the end of a branch only. The ooecial cell has the same
structure as the other cells and does not present the peculiarities found in Didymia
simplex.
S. Bugula dentata, Lamourou.x. Busk, Brit. Mus. Cat., I. 46.
The zooarium is delicate and of a leaden blue colour. There are two spines at
the upper inner angle, the lower one being the shorter ; one spine at the upper
outer angle, and two placed together further down on the outer side. All the spines
point upwards except the last on the outer side which is directed inwards.
9. Bugula avicularia, Linnaeus.
There is only one small fragment of this species.
10. Bugula yieritina, Linnaeus.
Growing on a Nautilus shell. A\dcularia occur at the base of a few cells.
11. Tubucellaria cereoides, Ellis and Solander.
Very common.
12. Cribrilina radiata, var. /3, Hincks. Hincks, Brit. Mar. Pol, p. 185.
The zoarium forms a silvery white crust over Terebratella shells. The zooecia are
oval and very minute (from '20 to -29 mm. in length x '14 to '20 mm. in width).
The ridges are not prominent; the interstitial pores are large. The central keel is
inconspicuous ; the small umbo below the mouth is most clearly seen in the fertile
cells. The ordinary zooecia bear six spines round the mouth, the fertile cells two on
each side. The ovicell has an indistinct keel. The avicularia occun'ing between the
cells are elongate ; vibraculoid appendages are wanting.
13. Memhranipora rudicifera, Hincks, var. intermedia, Kirkpatrick. Kirkpatrick,
Sci. Proc. Roy. Dub. Soc, Vol. VI. p. 615.
The form of the spines characteristic of Kirkpatrick's variety is clearly seen in
the young zooecia near the margin. On one side of the orifice is the .spine bearing
the avicularium, and the horizontal bifurcating branch sjoreading over the whole area.
This limb bifurcates repeatedly, the ultimate branches being slender. On the opposite
side there is a bifurcated spine of which the vertical branch is again bifurcated at
444 REPORT OX THE POLYZOA COLLECTED BY DR WILLEY
the tip ; the horizontal limb is also deeply bifurcated and each branch so formed
divides twice.
In some of the more central zooecia, some of the branches have met across the
area; the fusion has however not progressed so far as in the typical Hiantopora
ferox, MacGillivray, (Membranipora radicifera var. ferox, Kirkpatrick) so that here
we have an intermediate stage between the two varieties " intermedia " and "ferox."
15. Membranipora irregularis, d'Orbigny.
D'Orbigny, Voyage dans I'Amer. M^rid. viii. 56.
This is fairly abundant, growing round Echinid spines.
17. Membranipora punctigera, Hincks. Ann. Mag. Nat. Hist. 5. Vlll. 18S1, p. 4.
One colony growing on a Nautilus shell. No ooecia present.
19. Monoporella polymorpha, n. sp. PI. XLII., figs. 3 and 3 a.
Zoarium encrusting. Zooecia large, roughly hexagonal in shape, convex, rising
slightly towards the orifice, separated by distinct lines ; the surface in young cells
covered with large opaque dots, these being obscured later by the growth of a
brownish cuticular epitheca ; below the orifice a little brown patch.
Measurements of zooecia 1"07 mm. to 1"34 mm. in length x '86 mm. to 1'12 mm
in width.
Opercula calcareous, covered by a chitiuous membrane.
Opercula of two kinds :
(1) the form occurring most frequently; arched above with a straight lower
margin ; bordered by a dark line, the greater part of the operculum being coloured
brown leaving a light rim above.
Measurements : -3 to •34 mm. in width x -26 to '28 mm. in length.
(2) not quite so numerous as the first ; sides of the operculum sloping out-
wards from the straight lower margin ; upper border drawn out into three lobes ;
here also a semi-circular brown patch occupying the larger part of the operculum.
Measurements : Width at base of operculum from '3 to "So mm.
„ top „ „ '42 to '43 mm.
Length „ „ -32 to '39 mm.
In the bays between the lobes of the operculum and also on the sides just
below the lateral lobes are little circular scai's, showing the presence of spines absent
on zooecia bearing ordinary opercula. Rosette plates unusually large. Ooecium conical,
with well marked ridges and deep grooves alternately radiating from the apex ; from
four to six indentations on the floor of each groove. Operculum of the fertile cell
being of the more ordinary form, the upper margin arched, the lower straight or
slightly curved inwards.
Measurements of ooecium 96 mm. in width x -93 mm. in length.
This is a remarkable species being noticeable on account of its size and of the
FROM THE LOYALTY ISLES, NEW GUINEA AND NEW BRITAIN. 445
occurrence of two kinds of zooecia. The one kind possesses opercula of an ordinary
form and are without spines ; the other are characterised by large lobed opercula
and by the presence of spines in the bays between the lobes and on the sides of
the orifice. In one zooecium there is a slight undulation in the operculum below
the lateral spine so that this also lies in a bay ; this seems to suggest that the
bays may be due to a retardation of growth produced by the presence of spines.
No reason can at present be suggested for the existence of these two kinds of
opercula. The zooecia contain similar polypides ; those with the lobed operculum
possess slightly stronger opercular muscles.
20. Monoporella spinifera, n. sp. PI. XLII., fig. 4.
There are two small pieces.
Zooarium encrusting. Zooecia hexagonal, separated by depressed lines ; surface
covered with large poi-es ; the intervening parts studded with minute nodules ; orifice
horse-shoe shaped, surmounted by ten short brown spines.
Operculum calcareous, covered with a brown chitinous membrane ; horse-shoe
shaped within the orifice, the distal wall of the zooecium raised into a shelf projecting
into the cavity, the remaining space being closed in by a membranous diaphragm.
There is one peculiar zooecium in which the diaphragm is calcified, two small holes
being left for the opercular muscles. In this zooecium the orifice is drawn out into
lobes between the spines.
Ooecia ?
Measurements: zooecium l'2mm. in length xl'lSmm. in width; operculum ■27 mm.
in length x '32 mm. in width.
21. Schizoporella biaperta, Michelin. Hincks, Brit. ilar. Pol. p. 25-5.
Fairly abundant.
22. Schizoporella torquata, Quoy and Gaimard.
Only one fragment.
23. Schizoporella triangula, Hincks. PL XLII., fig. 5. Hincks, Ann. Mag. Nat. Hist.
(5) VIII. p. 12, 1881. Waters, Ann. Mag. Nat. Hist. (5) XX. 1887, p. 191.
The single specimen from New Britain seems on the whole to resemble Hincks'
species, though it dififers from it in some particulars. The surface is generally
reticulo-punctate without nodules, except in some old zooecia where small nodules are
found. The a\'icularia are placed on a small circular elevation from which lines pass
to the sides of the orifice marking out a distinct raised area. These avicularia are
not elongate but small and circular with a rounded mandible. However the form in
this species seems to var}' as in a specimen from Port Western, Melbourne, some
avicularia are elongated while those on adjacent zooecia are rounded. The ooecium
has a rough punctured surface, the older cells possessing also small nodules. The pro-
jecting processes and teeth on the margin of the ovicell described in Hincks' species
are absent.
27. Schizoporella depressa, n. sp. PI. XLII., fig. 6 and 6a.
Zoarium encrusting. Young zooecia ovate with a smooth surface; later the wall
at some little distance from the orifice becomes thickened so that eventually the
w. IV. 60
446 REPORT ON THE POLYZOA COLLECTED BY DR WILLEY
orifice lies in the centre of a hexagonal depression surrounded by the thickened
calcareous wall of the zooecium. Orifice arched above with a wide sinus below ; at
the junction of the two parts a minute denticle on each side.
On each side of the orifice in the young cell a minute oval avicularium placed
on a small elevation ; large vicarious avicularia with broad duck-bill shaped mandibles.
Ooecia ?
Measurements of zooecium : "29 mm. in width x "35 mm. in length.
32. Lepralia feegeensis, Busk. PI. XLIII., fig. 7. Busk, Challenger Report, xxx.
p. 144.
This species is abundantly represented in the collection. The opercula differ slightly
from those of the Challenger specimens. In the latter, the lower margins of the
opercula are straight or slightly curved inwards ; the Lifu specimens have the lower
margin slightly curved outwai-ds forming an indistinct sinus.
Ooecia are plentiful ; these do not occur on the Challenger specimens, and have,
I believe, not been described. The ooecium is globular and depressed, occupying the
greater part of the zooecium behind it. It is covered with large opaque granulations.
The fertile cells bear no a\'icularia.
Measurements of the ooecium : '64 to '74 mm. in width x "6 mm. in length.
33. Lepralia poissonii, Audouin.
There is only a tiny colony of this, growing over Lepralia tuberculata (n. sp.).
The vibracula are placed at the side of the orifice in line with the upper margin.
The vibraculoid mandibles are long, being equal to twice the length of the cell and
lie parallel with one another.
34. Lepralia tuberculata, n. sp. PI. XLIII., fig. 8.
Zoarium encrusting, in the form of a thick disc, the zooecia occurring only on
the upper face. Zooecia contiguous, arranged in radiating rows which become less
regular towards the centre ; cells rising towards the orifice giving the zoarium an
uneven surface ; surface of the individual zooecium smooth and shining ; above the
orifice eight marginal spines. Operculum orbicular or slightly coarctate with a rounded
lower border. On each zooecium placed at the sides of the orifice, two elongated
a\-icularia with blunt spatulate mandibles directed upwards ; numerous large vicarious
avicularia with duck-bill shaped mandibles. Ooecium globular, sub-cucullate, smooth,
reaching as far back as the orifice of the cell behind.
Measurements :
zooecium '3 to SB mm. in width x -32 to "42 mm. in length,
operculum '1 mm. x ■! mm.
ooecium "18 mm. in width x "lo mm. in length.
There is only one colony of this species.
35. Lepralia calyciformis, n. sp. PI. XLIII., figs. 9 and 9 a.
Zoarium unilaminar and cup-shaped, being attached at the base ; zooecia occurring
FROM THE LOYALTY ISLES, NEW GUINEA AND NEW BRITAIN. 447
in rows radiating from the centre and being separated by depressed lines marked by
small indentations; surface smooth and shining. Orifice sub-central, surmounted by
eight spines. Operculum coarctate with an arched upper margin and a slightly convex
lower margin. Placed far back on each side of the orifice a minute avicularium on
a small elevation, mandible rounded, and directed inwards and upwards. Distributed
over the zoarium, other minute oval avicularia with rounded mandibles.
Ooecia ?
Measurements : zooecium •82 mm. in width x "SI mm. in length, operculum "08 mm.
in width x '11 mm. in length.
This species which is here represented by a single colony also occurs in the collec-
tion from ToiTCs Straits which Dr Haddon has presented to the Cambridge University
Museum of Zoology. It l)ears a superficial resemblance to Lepralia tuber culata, the
species described above, but is distinguished from it by the shape of the zoarium,
the size and shape of the opercula and the form of the avicularia.
36. Smittia marmorea, Hincks. PI. XLIII., fig. 10. Hincks, Brit. Mar. Pol., p. 350.
The collection contains one small fragment which seems to correspond with
Hincks' species. The sinus in the lower margin of the secondary orifice is broad and
the notches on each side are very prominent. The pores at the edge are very large
.and the avicularia are longer than in Hincks' species, extending the whole length of
the zooecium.
No ooecia are seen on this fragment.
37. Mucronella articulata, n. sp. PI. XLIII., fig. 11.
Zoarium disc-shaped and encrusting. Zooecia arranged in lines, the cells being
indistinctly separated from one another ; the surface finely granular. Orifice orbicular
and below it a small mucro ; round the orifice eight long jointed spines standing up
vertically ; each spine consisting of three joints, the terminal segment ending in an
expanded funnel. Placed high up, one on each side of the orifice in the younger
zooecia, two minute avicularia with rounded mandible dii-ected inwards. Distributed
over the zoarium, large vicarious immersed avicularia with broad duck-bill shaped
mandibles.
Ooecia ?
Measurements : zooecium "42 mm. in width x 26 mm. in length, operculum
•1 mm. X "1 mm.
The characteristic feature of this species is the presence of jointed spines with
the terminal segment expanded into a funnel. In most of the central zooecia only
the two lateral spines have this cup-like termination, the remaining six being shorter
and possessing ends which are not expanded but often pointed. However the fact that
most of the marginal zooecia possess eight spines with expanded ends suggests that
the others have lost their terminal joint.
There are three colonies of this species.
60—2
448 REPORT ON THE POLYZOA COLLECTED BY DR WILLEY
38. Rhynchozoon bispinosum, Johnston. Hincks, Brit. Mar. Pol. p. 38.5.
The mucro is small and no spines are visible even on the marginal cells.
39. Rhynchozoon crenulatum, Waters. Rhynchopora crenulata, Waters, Ann. Mag.
Nat. Hist., 5, xx. 1887, 195.
There is only one colony of this species.
40. Escharoides spinigera, n. sp. PI. XLIIL, fig. 12.
One small encrusting colony.
Zooecia ovate, surface smooth and polished. Primary orifice sub-orbicular with
an entire margin ; above the orifice eight spines ; the secondary orifice with a sinus
on one side of the lower margin ; on the inner side of the peristome directed
towards the sinus a minute avicularium with a bluntly pointed mandible ; on some
zooecia, placed by the side of the orifice, an avicularium with a spatulate mandible
directed upwards.
Ooecia ?
Measurements : zooecium from "29 to '37 mm. in width x '30 to '43 mm. in length,
operculum "06 mm. x "08 mm.
The avicularium on the inner side of the peristome is visible in only a few cells.
41. Gellepora simplex, MacGilli\Tay. McCoy. Prodr. Zool. Vict. p. 241, pi. 165.
A common species in Lifu.
44. Cellepora longirostris, MacGillivray. Pi. XLIII., figs. 15, 15 a. MacGilL, Trans.
Roy. Soc. Vict. 1884, p. 113.
The collection contains two fragments which appear to belong to this species.
Both are encrusting, but from the centre of one there projects a short process which
suggests that the specimen may be simply the encrusting base of an erect colony —
the erect habit being characteristic of Cellepora longirostris. The young zooecial
stages described by MacGillivray are clearly seen on the margin of the specimens.
The orifice has a rounded sinus, and on one side is the small avicularium which is
later concealed by the growth of the peristome. The opposite edges of the latter
arch over the sinus and form a rounded opening which is subsequently filled in.
Finally there grows up from the sub-oral part of the peristome an enormously developed
rostrum, the free apical part of which is, as a rule, coarsely serrated. The rostrum
has a thick base, and generally bears on its outer face an elongated avicularium
directed downwards. The unusual size of the rostrum is a striking characteristic of
the Lifu specimens. The ovicell is globular, and is almost concealed by the peristome.
The front is sculptured in a radiate manner.
45. Retepora phoenicea, Busk. Busk, Brit. Mus. Cat. ii. 94. Busk, Challenger
Report, XXX. 124.
This species is plentiful in. Lifu.
FROM THE LOYALTY ISLES, NEW GTJINEA AND NEW BRITAIN. 449
46. Retepora denticulata, Busk. PI. XLIIL, fig. 13. Busk, Challenger RejDort,
XXX. 109.
The dorsal avicularia seem to be more numerous than on the Challenger specimens.
They occur laterally on the dorsal surface and on the trabeculae connecting the branches.
The beak is bluntly j^ointed and is bent up at the end ; the mandible is triangular
with a broad base ; the pointed apex is bent down. The ooecia in many respects
resemble those of Retepora apicidata (Busk, Challenger Report, XXX. 108). They are
globular, smooth and sub-cucullate. There is a median grooved keel on each side of
which there is a slight longitudinal depression. The upper lip of the opening is trifid.
47. Crisia acideata, Hassall. Hincks, Brit. Mar. Pol. p. 421. Hassall, Ann.
Mag. Nat. Hist. (.5) vii. 368. Harmer, Q.J. M.S. 18.91, p. 132.
There is one small colony growing on a sea-weed. In the absence of an ovicell,
it may provisionally be referred to this species.
48. Crisia denticulata, Lamk. Hincks, Brit. Mar. Pol., p. 422.
There are several colonies growing on a large crab. No ovicells are present on
the specimens.
•50. Idmonea interjuncta, MacGillivray. MacGill. Trans. Roy. Soc. Vict. 1885, p. 137.
Common.
.51. Idmonea radians, Lamarck. Busk, Brit. Mus. Cat. iii. 11.
Very common.
.52. Idmonea australis, MacGillivray. McCoy, Prodr. Zool. Vict. T. PI. 68, p. 30.
Several rather young colonies.
53. Entalophora delicatida, Busk.
Pustulopora delicatula. Busk, Brit. Mus. Cat. iii. 20.
Several small colonies.
54. Hornera spinigera, Kirkpatrick. Kirkpatr., Ann. Mag. Nat. Hist. 6, i. 1888, 83.
One large fine colony.
59. Lichenopora californica, D'Orb.
Discoporella californica. Busk. Brit. Mus. Cat. ill. p. 32.
One small colony growing on a Nautilus shell.
60. Lichenopora truncata, n. sp. PI. XLIII., figs. 14 and 14 a.
Zoarium convex, flattened on the top ; shaped like a truncated cone, bordered by
a thin lamina ; central flattened area occupied by cancelli. Zooecia ai-ranged in long
uniserial rows of almost equal length, there being about fifteen zooecia in a mw in
the type specimen. Zooecia not raised above the surface. Orifice circular, the peri-
stome being produced on the upper side; margin of the peristome entire or slightly
450 REPORT ON THE POLYZOA COLLECTED BY DR WILLEY
produced at the angles ; between the zooecial rows four series of cancelli which are
not denticulate ; those belonging to the two central rows of each interspace as large
as the orifices of the zooecia, the lateral cancelli being smaller.
The colony is about 5 mm. in diameter. Two specimens only occur in the collection.
61. Radiopora cristata, Busk. Busk, Brit. Mus. Cat., III. 35.
There are numerous fine colonies of this species.
62. Fasciculipora carinata, Ortmann. Ortmann, "Jap. Bryo. Fauna." Arch. f.
Naturg., 56, i. 1890, p. 65.
One colony.
63. Zoobotryon pelliicidum, Ehrenberg. Reichert, Abh. k. Akad. Berlin, 1869. Ii.
These specimens differ slightly from the Mediterranean species in the arrange-
ment of the zooecia. Here the individuals are more crowded at the ends of the
branches, while there are longer spaces devoid of zooecia on the main branch.
Otherwise the characters seem to be those of Ehrenberg's species.
EXPLANATION OF PLATES XLII AND XLIII.
All the figures were drawn by means of a camera lucida; figs. 1, 5, 6, 8, 9, 10, 11,
12 and 13 with a Crouch objective A with front lens on; figs. 2 and 2 a, 3 and 3 a, i,
7 and 14 with Crouch objective A ^vith front lens off. All the figures were then reduced
by one half.
Fig. 1. Canda retiformis, Pourtales.
Fig. 2 and 2 a. Didymia triserialis, n. sp.
Fig. 3 and 3 a. MonoporeUa polymorpha, n. sp.
Fig. 4. MonoporeUa spinifera, n. sp.
Fig. 5. Schizoporella triangula, Hincks.
Fig. 6. Schizoporella depressa, n. sp.
,, 6 a. Young zooecia of same.
Fig. 7. Lepralia feegeeusis, Busk, showing ovicell.
Fig. 8. Lepralia tuberculata, n. sp. ,
Fig. 9. Lepralia calyciformis, n. sp.
,, 9 a. Colony of same, four times natural size.
Fig. 10. Smittia marmorea, Hincks.
Fig. 11. Mucronella articulata, n. sp.
Fig. 12. Escharoides spinigera, n. sp.
Fig. 13. Ovicells of Retepora denticulata, Busk.
Fig. 14. Lichenopora truncata, n. sp. Part of a colony seen from the side.
„ 14 a. Colony of same, four times natural size.
Fig. 15. Cellepora longirostris, MacGill. The rostrum below the ovicell has been broken off.
„ 15a. Young zooecia of same.
._EY. AOO
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PHILIPPS. POLYZO...
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THE HYDROID ZOOPHYTES COLLECTED BY DR WILLEY IN
THE SOUTHERN SEAS.
By LAURA ROSCOE THORNELY,
University College, Liverpool.
With Plate XLIV.
The Hydroids in this collection, though many of them are broken from their
colonies in a tantalizing way, were evidently so full of life when gathered, and are
so beautifully preserved, that to the smallest details they have been interesting to
examine. Of two or three species, as I have noted uuder the descriptions of them,
there are considerable quantities; of others, few, but complete, colonies; and of
several too little to allow me to do more than identify the genus to which they
belong. Of these last there is a Gryptolaria, probably C. pulchella Allman, a
Eudendrium, a Lafoea, two Thuiarians, an Ophiodes, and a Sertularella, probably
S. angulosa.
I am much indebted to Dr Willey for trusting me with his specimens, and I should
like to offer my very best thanks to Professor Herdman, D.Sc, F.R.S., for his very
great help with advice and with books.
Family. Atractylid^, Hincks.
Genus. Hydranthea, Hincks.
Hydranthea australis, n. sp.
There are several little colonies of this species creeping over other zoophytes.
The stem is very short, enclosed at the base in little cup-like extensions of the
polypary, as in Hydranthea margarica, Hincks', which also has numerous tentacles
in a single verticil and large gonophores like this species. The tubercles of thread
cells at the base of the tentacles, and the four branched vessels in the gonophore,
characteristics of Hydranthea, are not visible in these specimens, but the whole appear-
ance points to the species being of this genus and closely related to H. vmrgarica.
1 British Ilydroid Zoophytes, 1808, p. 100.
452 HYDROID ZOOPHYTES COLLECTED BY DR WILLEY IN THE SOUTHERX SEAS.
The position of the gonophore, rising from the side of the cup of the hydrotheca, instead
of from the stolon and within a cup of its own, is the only marked difference which
makes it necessary to separate the species.
Locality. Blanche Bay, New Britain ; growing among other zooph}i:es attached
to ropes and ti.sh-ba.skets down to 40 fathoms.
Gexus. Bougainvillia, Lesson.
Bougainvillia muscus Allman, PI. XLIV. Figs. 2, 2 a, 2 b.
There are colonies of this Bougainvillia covering the cast spines of a Cidaris. They
are taller than the description of B. muscus, as described by Hincks', corresponding
thus more with his form intermediate between that species and B. rainosa, but they have
not the compound stem of this form though they have the sanded polyparj-.
These colonies have gonophores (Fig. 2 h) borne on the stem, some way below the
polypites, two or three together on a branched offshoot.
From a separate collection taken from ropes and fish-baskets and floats there
are also some colonies which are shorter and have curious tendrils, some simple, some
branched (Fig. 2 a), and of varjang lengths, attached to the stems below the poh'pites.
The characters in both these sets of specimens do not seem sufficiently marked to
justify the formation of a species apart from Bougainvillia muscus Allman.
Locality. Blanche Bay, New Britain.
Family. Tubulakiid^, Hincks.
Genus. Ectopleura, L. Agassiz.
Ectopleura pacifica, n. sp., PI. XLIV. Figs. 1, 1 a.
Of this fine species there are a profusion of colonies, detached from floats, fish-
baskets, &c. The stems are simple, unringed, diminishing in width towards the base
and about 2 cm. high, rising from a creeping stolon which connects the colony. There
are some processes given off fiom the stem near the base in many of the specimens
which probably help to attach the colonies to the objects thej" gi-ow upon. The
polypite is abruptly marked off from the supporting stalk. The tentacles are disposed
in two verticils of from 18 — 20 each. The distal tentacles appear to be not wholly
filiform (see Fig. 1), but this may be the effect of contraction of the preserved specimen.
The gonophores are borne in clusters of from seven to nine on branched peduncles
which spring from the body of the polypite, between the two sets of tentacles. The
most highly developed among them have two long tentacles (Fig. 1 a), and there are
indications of what may be the eight longitudinal ribs formed of the linear series of
thread cells found in the genus Ectopleura. The above species is quite distinct from
the only other Ectopleura, E. dumortierii. Van Beneden.
Locality. Blanche Bay, New Britain.
' British Hydroid Zoophytes, 1868, p. 111.
HYDROID ZOOPHYTES COLLECTED BY DR WILLEY IN THE SOUTHERN SEAS. 453
Family. Campanulariiuae, Hincks.
Genus. Obelia, Peron et Lesueur.
Obelia linearis, n. sp., PI. XLIV. Fig. 6.
There are only two little rooted specimens of this species, 7 mm. in height. The
stem is simple, horn-coloured below, transparent above, and is branched (see Fig. G).
The h3-drothecae are deep, their margins have about twelve blunted teeth around
them. There are usually longitudinal lines to be seen on the hydrothecae, the result
probably of the collapsing of their delicate sides.
The gonothecae are borne in the axils of the peduncles, they are elongate oval,
smooth, and with a prominent rim to the orifice.
The gonozooids seen within the gonothecae show signs of tentacles in those that
are uppermost in the gonothecae.
Locality. Blanche Bay, New Britain.
Obelia delicatula, n. sp., PI. XLIV. Fig. 7.
This is a smaller, more delicate form than the last, and unbrauched, but the
hydrothecae and gonothecae are much the same in their characters.
The stem reaches 4 mm. in height, borne on a creeping stolon, which carries the
gonothecae also.
The stems are ringed at their bases, and here and there above, and are also several
times ringed under the hydrothecae (Fig. 7). The whole colony lias much the appear-
ance of Campamdariu phi/coci/uthus, Allman', but the zooids in the gonothecae show the
budding tentacles of the free swimming medusoid, showing that it does not belong to
that genus.
Locality. Blanche Bay, New Britain, 40 fathoms.
Obelia serrulata, Bale (sp.), PI. XLIV. Fig. 5.
The large.st piece among the fragments collected of this species is a little over 1 cm.
in height. It is branched and has a strong compound stem formed by the downward
growth of stolons from the peduncles of the hydrothecae.
The hydrothecae are very broad above, tapering downwards towards the base. The
floor is raised above the base so as to leave a large cavity beneath it, and this
character, together with that of the marginal teeth, agrees with Mr Bales'- description
of Gampanularia ? serrulata, Bale, found at Port Jackson. His specimens were delicate,
and there being no gonothecae present makes it appear likely that they were immature.
The gonotheca of the present species is oval, truncated above, and borne on the
stem, and contains zooids with budding tentacles (Fig. 5).
Locality. Blanche Bay, New Britain : from ropes and fish-baskets in 40 fathoms.
1 Voyage of H.M.S. CliaUeuger, Report on llie Ilijtlroiiin, Pt ii.
'•' Proc. Linn. Soc. Neic South U'ulex, Vol. in. 1888, p. 7J7.
w. IV. 61
454 HYDROID ZOOPHYTES COLLECTED BY DR WILLEY IX THE SOUTHERX SEAS.
Genus. Campamdaria, Lamoiiroux (in part).
Campanularia hrevithecata, n. sp., PL XLIV. Fig. 8, 80, 8 6.
Thi.s form has a simple, unbranched stem, a little more than 1 cm. in length,
very slender, with several rings at the base (Fig. 8 a) and one or two at intervals
up the stem, while directly below the hydrotheca there is one spherical ring as in
Campanularia volubilis, Linnaeus.
The hydrotheca is extremely short, so as only to cover about one-third of the
polypite when it is extended, and has an even rim (Fig. 8).
The polypite has about twenty tentacles, and a remarkably large trumpet-shaped
proboscis, suggestive of a Eudendrium from which the species is widely separated by
its other characters.
The gonothecae (Fig. 8 h) present rise from the stolon on short pedicels, and
have the appearance of those of Campamdaria calicidata, Hincks, as figured by Hincks^
and contain one or two large sporcsaes with four gastro-vascular canals to each.
C. calicidata Hincks, as figured by Mr Bale^ has much the appearance of my
specimen, but without the polji^ite the proportional sizes of polypite and calycle are
not obvious. In some of ilr Bale's specimens from Port Jackson the calycles have not
the thickened wall usual in C. calicidata, which would bring it nearer to our form, but
there are still sufficient grounds for considering this a new species on account of the
smallness of the theca and the largeness of the polypite proboscis.
Locality. Blanche Bay, New Britain, one colony covering the body of a Lepas,
others detached from ropes and fish-baskets.
Genus. Gonothyraea, Allman.
Gonothyraea longicyatha, n. sp., PI. XLIV. Fig. 4, 4 a.
There are a large number of complete colonies of this strikingly pretty form.
Their height is about 3 cm., they are much branched, the main stem is deep brown
in colour, and offshoots from the bases of the pedicels of the hydrotheca, growing
downwards, form a strong compound stem (Fig. 4 0), as described by Dr Hartlaub^ for
Obelaria gelatinosa, and by Mr Bale'' for Obelia ? spinulosa.
The hydrothecae are remarkably long and narrow, and the castellations on their rims
so deeply grooved as to make sharp, needle-like points of their edges (see Fig. 4).
The gonothecae are very numerous, borne on short ringed pedicels in the axils
of the branches. The}' are oval and truncated at the top. In nearly all cases they are
flattened laterally, whicli makes the top fall in, forming a groove, but this is probably
a result of the pi-eservation of a delicate form.
1 British Hydroid Zoophytes, 1868, PI. 31, Fig. 2 d.
- Proc. Linn. Soc. New South Wales, Vol. 111. 1888, p. 755.
■■' Jleeresfauna von Helgoland 1897, Zweiter Bericht, Kiel.
■* Proc. Linn. Soc. New South }]'ales'. Vol. iii. p. 756.
HYDROID ZOOPHYTES COLLECTED BY DR WILLEY IN THE SOUTHERN SEAS. 455
I have only seen one external capsule, and that is not perfect, but unfolding tentacles
can be seen on the uppermost sporosac in several of the gonothecae.
This description agi-ees with that of Gonothyraea hyalina, Hincks', to some extent,
but here there is the fascicled stem, and the hydrothecae, as well as the points on
their turrets, are moie drawn out, while their total size is less. If what appears to be
an external capsule is in reality an escaping medusiform zooid, the species may be
an Ohelia.
Ohelia longicyatha, AUman-, is like the present species in the shape of the hydrothecae,
but not in that of the teeth on their margins. Campanulana ? spinulosa, Bale^ agrees
in the shape of the hydrotheca and its teeth and in the polysiphonic stem, and as
gonothecae have not been found for this form it is possibly this species. From the
description by Mr Clarke of Ohelia hidentata* it is evidently a much larger species
than this.
LOCALITV. Blanche Bay, New Britain : from floats and ropes attached to fish-
baskets, some from a depth of .50 fathoms.
Family. Sertulariidae, Hincks.
Genus. Sertularia, Linn^ (in part).
Sertularia pusilla, n. sp.
This is a minute species, only 3 mm. in height, and looking in all ways
only half the size of the Pasythea quadridentata, Ellis and Sol., amongst which it is
growing.
The opposite hydrothecae join each other in front and are separated behind.
They are very long and slender, and delicate looking, as are the internodes which
bear these one pair on each. The polypites show almost black through the transparent
polypary.
Locality. Lifu, Loyalty Islands.
Sertulana littoralis, u. sp.
There are several specimens of a little Sertularia of a bright brown colour, looking
like S. pumila, Linnaeus. They are smaller than that form, however, being only 7 mm.
in height, and with the branches in the one specimen that has any, alternate instead
of opposite. These are placed on the internodes, below the hydrothecae of three
successive internodes, beginning on the third from the base. The hydrothecae are
> British Hydroid Zoophytes, 1868, p. 184.
2 -'Report on the liydroida collected during the exploration of the Gulf Stream," by L. F. de PourtalSs.
Memoirs Museum of Comparative Zooloyy, Harvard University, Vol. v. (1877), p. 10.
= Proc. Linn. Sac. ^'eu■ South Wales, Vol. in. 1888, p. 756.
•• "Description of new and rare Hydroids from the New England coast." Transactions of the Connecticut
Academy of Arts and Sciences, Vol. in. Pt. 1.
61—2
45fi HYDEOID ZOOPHYTES COLLECTED BY DR WILLEY IN THE SOUTHERN SEAS.
opposite in all cases, and there are none in the axils of the branches. They are in
contact with each other in front, separated behind, and have a horizontal fold crossing
the cell as in S. loculosa, Busk.
There are no gonothecae.
Locality. Lifu, Loyalty Islands; littoral.
Gexus. Pasythea, Lamouroux.
Pasythea quadridentata, Ellis and Solander. Nat. Hist, of Zoophytes, London, 1786.
There are a few fragments of this species.
Locality. Lifu, Loyalty Islands.
Family. Plujil'lariidae, Hincks.
Genus. Aglaophenia, Lamouroux (in part).
Aglaophenia hellis, n. sp.
This -species has almost the exact appearance of Professor AUman's' figure of
Aglaophenia Macgillivrayi, Busk (sp.), and most of its details correspond with his
description of them, but there are a few marked differences which I think make a
separation necessary.
There is the fascicled stem with opposite pinnae as in A. Macgillivrayi, gi\'ing
off secondary alternate pinnae which carry the hydrothecae, and there are branches
taking the places of pinnae at irregular intervals and resembling the stem in structure.
It is remarkable in this species the thickness of even the pinnae, the hydrothecae
bearing pinnae, fomiing merely a feathered line up the middle of a thick " woody "
stem, which is .spotted by the lateral communications between its tubes spoken of by
Professor Allman" as a characteristic of this gi-oup.
The hydi'othecae have the same intrathecal ridge as m A. Macgillivrayi, the
mesial sarcothecae are adnate for their whole length, the same height as the hydro-
thecae, bithalaraic and canaliculate, and the lateral sarcothecae correspond, but I
cannot see the intracauline ridges, and the margins of the hydrothecae have a little
projecting point on either side.
The corbula is closed, with about eight series of costae, but there is no spur-like
sarcotheca at the base, nor cup at the summit of each costa which give so striking
an appearance to Professor Allman's figure of the corbula in A. Macgillivrayi.
Viewed from the front my corbulae show that the ribs are disjointed in the
middle, the two halves rise from different points at their bases and do not therefore
meet above. This form would agree with Professor Allman's^ idea that these corbulae
1 Voyage of the 'Rattlesnake,' Vol. i. 1852, p. 400.
- Voyage of H.M.S. Challenger, Pt 1, 1883, p. 5.
3 Ibid., Pt. I. p. 11.
HYDROID ZOOPHYTES COLLECTED BY DR WILLEY IN THE SOUTHERN SEAS. 457
are formed of the mesial sarcothecae of a hydrocladium or ultimate pinna, wliich, deprivi^d
of their hydrothecae and lateral sarcothecae, are thrown off to left and right alternately
up the pinna, giving off secondary sarcothecae which continue the ribs upwards. The
position of the corbula taking the place of a hydrotheca bearing pinna and the
number of the costae corresponding with the number of hydrothecae on these pinnae,
allowing for the one complete hydrotheca which is always present below the corbula
in these specimens, makes it seem natural that the corbulae are formed thus.
There are only a few species known of Aglavphenia which have the doubly
pinnate ramification and the bithalamic form of mesial .sarcotheca, and A. Macr/illivrai/i
is the only representative in Australian seas.
Locality. Engineer group, British New Guinea.
Gexus. Pluinularia, Lamarck (in part).
Plumidaria compactu, n. sp., PI. XLIV. Fig. 3.
A good large quantity of this species was found. The colonies resemble the
description by Mr Bale' of a small variety of Plumularia setaceoides, Bale-. They
are pale horn colour, unbranched, 1'.5 cm. in height. The pinnae are alternate and
bear four hydrothecae at most, which have even rims and a mesial and two lateral
sarcothecae, while there is also a sarcotheca on an internode between every two hydm-
theca-bearing internodes and one on the lower part of each stem internode.
The gonothecae grow near the bases of the .stems, below the pinna-bearing
internodes. They differ from those of P. setitceoides, which are obliquely truncated a
little above the broadest part, in tapering upwards from the broadest part about as
much as they taper downwards below it and in terminating with a neat rim. These
gonothecae show clearly that this is not Mr Bale's P. setaceoides: it may be what
he described as a smaller variety, in which he found no gonothecae, but in that case
it must become the independent species — which I have described above as P. comjxicta.
Locality. New Caledonia.
' Catalogue of the Australian Hydroids, 1SS4.
^ JouTH. Mic. Soc. Vict., 11. (fig.).
EXPLANATION OF PLATE XLIV.
Fig. L Ectopleura pacijica, n. sp.
„ 1 a. The gonophores.
Fig. 2. Bougainvillia muscus, Allman.
„ 2 a. Tendrils.
,, 2 6. Gonophore.
Fig. 3. Plumularia compacta, n. sp.
Fig. 4. Gonotliyraea longicyatha, n. sp.
4 a. The stem showins the downward growth of the stolon.
Fig.
5.
Ohelia serntlata, Bale (sp.)
Fig.
6.
„ linearis, n. sp.
Fig.
7.
„ delicatula, n. sp.
Fig.
8.
Campanularia brevithecaia, n. sp.
J)
8 a.
Base of the stem.
»)
8 6.
The gonotheca.
WilIjEY Zoological, P.esul.ts
Plate XLIV.
Wcirt.Nrwmim litii.
THOKNEliY HYDKOiDS.
ASTROSCLERA WILLEYANA,
THE TYPE OF A NEW FAMILY OF SPONGES.
By J. J. LISTER, M.A., F.Z.S.,
Fellow of St John's College and Demonstrator in Comparative Anatomy in the
University of Cambridge.
With Plates XLV— XLVIII, and three Figures in the text.
CONTENTS.
Introductory Remarks.
E.XTERNAL CHARAeTERS, p. 460 :
of the Lifu .specimens ; of the Funafuti specimen.
Methods, p. 462.
The Skeleton, p. 463 :
microscopic characters ; mineralogical characters ; organic basis ; origin of the skeletal elements ;
the arrangement of the skeletal canals.
Mode or Growth, p. 466.
The Specific Relationship of the Specimens from the two localities, p. 468.
The Soft Tissues, p. 468 :
the gelatinous layer at the surface and the pores; the ciliated chamlicrs: the canal sj'stem.
Reproduction, p. 470.
State of Preservation, p. 470.
Affinities, p. 471.
Discussion of resemUances to Madreporaria, p. 471 ; the isolated position of Astrosclera among
sponges, p. 472 ; comparison with the Pharetrones, p. 473 ; on the Spherulitic structure seen
in the Pharetrones from St Oassian and elsewhere, p. 476.
Definition of the Family Astroscleridae (new family, genus and species), p. 479.
Conclusion, p. 480.
In the collections brought home by Dr Wille}- from the \\'(>slern Pacific were
four specinien.s of a peculiar hard white oigauism which lie found growing on dead
coral, at a depth of 3.5 fathom.s, in Sandal Bay, Lifu, Loyalty Islands. These he
placed in my hand.s for examination. A .short account of the results I had arrived
at was given at the meeting of the British Association at Dover in September, 1899.
As the organism appeared to be new, the name Astrosclera willeyuna wa.s proposed
460 ASTROSCLERA WILLEYANA, THE TYPE OF A yE\V FAMILY OF SPONGES.
for it'. In November I received from Mr Kirkpatrick of the British Museum another
specimen of the same or a closelj" allied organism, which was collected in the boring
expedition to Funafuti, one of the Ellice Islands h'ing about a thousand miles to the
N.E. of Lifu. This was obtained at the depth of 100 fathoms on the outer slope of the
western side of the island.
Before proceeding to details it may be stated that the hard parts consist of solid
polyhedral elements united to form a continuous skeleton. This is penetrated by canals
which branch and anastomose freely, and open to the exterior by apertures on the upper
surface. The canals are occupied by soft tissues which also cover the upper surface,
and are continued for some distance down the sides.
External Characters. The Lifu specimens are cj-lindrical in shape, and measure
about 10 mm. in height and 5 mm. in breadth. The base is slightly spreading, the
sides smooth and imperforate, and faintly marked by annular constrictions, and the upper
surface gently convex. In three of the specimens the growth has occurred at right
angles to the surface of attachment (Fig. 1), but in the fourth (Fig. 2), whose attach-
ment appears to have been to a vertical surface, the axis is curved. The rounded upper
surface is closely pitted by the openings of the canal system, those at the periphery
being smaller than the others.
In three of the specimens the openings are disposed indefinitely (Fig. 2 «), but in
the fourth (Fig. 1 «) there is a somewhat different arrangement. The upper surface
of this specimen is oval in outline, and about a point nearer one end of the long
axis than the other some seven grooves are disposed in a radiate manner. The
grooves show a tendency to branch at their outer ends, before they are lost in the
irregularities of the surface ; and one of them, which lies in the long axis, gives rise
to two subsidiary grooves which however are not disposed symmetrically about it.
The openings of the canal system are to be seen in the floors of the grooves as well
as on the parts of the upper surface lying between them. The soft tissues here
cover the surface, and it is not obvious that the pores in the grooves, or at the
centre from which they radiate, are larger than the others. A comparison of it with the
Funafuti specimen, however, renders the existence of larger pores in these situations
not improbable. The grooves are probably the initial stages of radially disposed canals.
The specimen from Funafuti (Fig. A, 1 — .5) has grown attached by a short stalk
(A, 1, St.) about 6 mm. wide at the base, which expands into a broad, nearly circular
disc (d — d), convex above, and resembling the pileus of a mushroom in shape. The
diameter of the disc is about 20 mm., and the distance from the broken end of the
stalk to the centre of the upper surface of the disc is 1(J mm.
A smooth, imperforated, cortical layer covers the outer surface of the stalk and
under surface of the disc, and presents concentric ridges and grooves, marking lines
of growth (A, 2). The upper surface of the disc is in part perforated by pores, whose
arrangement is described below.
There appears to have been an interruption to the growth of the specimen after
the formation of the disc, and the later extension has taken place not uniformly, but
' The definition of the species, i.e. of the family which it constitutes, is here given for the first time, p. 479.
ASTROSCLERA WILLEYAXA, THE TYPE OF A NEW FAMILY OF SPONGES.
461
only from parts of its upper surface. At these parts rounded bosses of different sizes
have been formed (Fig. A, 1, and 2, b, b), and they, like the original disc, are perforated
by pores on their conve.x ujjper surfaces and covered on the sides by an imperforate
^liv ">"^
3. ^^^®2l^^'
■W
"-::k
Fig. a. The Funafuti Specimen of Astbosclera.
b, h. The bosses formed by later growth. ;/, ((. The disc-shaped expansion of the original growth. .<(. The
stalk of attachment.
1. Side view.
2. View from the base, with the stalk turned towards the spectator. Part of the specimen has been
cut away, and the base has been excavated by some boring organism.
3. View of the surface of the large bosses formed by later growth.
4. One of the radiate systems of large pores more highly magnified.
5. View of the cut surface of the original growth, showing the radiate arrangement of the large canals
in the interior.
cortical layer, which in some places appears to have spread from their base over the
adjoining regions of the surface of the original disc.
The growth of the bosses repeats the character of that of the original disc, expand-
ing from the base ; and the greater mass of the new growth appears to have been
formed by the fusion of three originally distinct bosses.
w. IV. 62
462 ASTROSCLERA WILLEYANA, THE TYPE OF A NEW FAMILY OF SPONGES.
It is further to be observed that the trend of growth of the later formed bosses
is in a direction deflected at an angle of about 60° from the axis of the original
growth. We may conjecture that this result has been produced by the shifting of
the position of the object — block of coral or what-not — to which the specimen was
attached.
The upper surface, whether of the original disc or of the later formed bosses, is
perforated by closely set pores (Fig. A, 3 and 4). These are sometimes isolated, but
often they open into curving and branching grooves, recalling those of the coral
Maeandrina. At seven places on the surface of the later growth, and at one on the
original growth, the lines of pores are seen to be disposed in a radiating manner
about so many centres. The lines about a centre vary in number from five to ten,
and at their outer ends they become lost among the smaller pores round about.
The pores along these lines are large (Fig. A, 4), being at least twice the diameter
of those distributed elsewhere over the surface, and a group of such pores is situated at
each of the centres from which the lines radiate.
There is then a differentiation of the pores, and hence of the canals, whose openings
they are, into two categories, viz. large ones disposed in radiating systems, and small
ones distributed between the systems. As will be seen later, the distribution of the
pores at the surface is the expression of an arrangement of the canals found throughout
the interior.
The soft parts proper to the organism were not present in the portions of this
specimen that I have decalcified, and a boring sponge, with characteristic pin-shaped
siliceous spicules and bundles of raphides, appears in the sections. Moreover, the surface
even of the most recently formed part is dotted over with the pink growths of the
Forarainiferan Polytrema, a brown alga has established itself at one point (these have
been omitted in the figures), and the base has been much excavated by boring organisms.
It appears, therefore, that this specimen of Astrosclera had died some time before
it was collected.
Methods. When collected by Dr Willey the Lifu specimens were put into strong
alcohol (over 70 °/„). In examining them two have been decalcified by means of dilute
acetic acid, added drop by drop to a dish of 70 °/^ alcohol containing them, until bubbles
of gas appeared. When decalcification was completed, the soft organic body was divided
longitudinally, and the parts embedded in paraffin and cut into sections — one half
into longitudinal sections, the other into transverse. Another specimen was dealt with
as follows. It was first divided longitudinally by a fret-saw, to allow the reagents to
penetrate the interior more readily. Some idea of the degree of hardness of the
skeleton may be gained from the fact that the lines of the sawing were sharply
marked on the cut surfaces. On the other hand, it was not so hard as to perceptibly
blunt the saw, a result which was soon effected by the skeleton of a specimen of
Polytrema, on which I made a preliminary experiment. The upper part of one half
of the specimen was sawn off", decalcified, and cut into sections, for comparison with
the sections of the other specimens. The remaining parts were then stained in borax
carmine for two days, passed through 90 °/„ and absolute alcohol into chloroform, and
ASTROSCLERA WILLEYANA, THE TYPE OF A NEW FAMILY OF SPONGES. 463
then placed in an open dish in a thin solution of gum copal in chloroform. After
some days, when the solution had become thick, the specimens were put on slides,
with the thickened gum about them, and placed on the shelf of the warm water bath
(60° C.) to harden. When the gum was hard, slices of them were cut with the
fret-saw, one half being cut longitudinally, the other transversely. Each furnished
three slices.
One surface of the slice was ground down smooth on a hone, and it was then
cemented to a slide with Canada balsam, with this surface downwards. When the
balsam was hard, the other surface of the slice was ground down until the section
was so thin that the edge began to break. It was then dried and covered with
Canada balsam and a coverslip.
This method is that of von Koch, as given in Bolles Lee's Vade-mecum. It
may be worth while to mention that in grinding the sections, lumps of ice were kept
on the hone in order that the copal (which is soft at GO" C.) might be a.s hard as
possible.
The structure of the hard and soft parts is well displayed in these preparations.
The fourth Lifu specimen I have kept intact.
Slices both vertical and tangential to the most recently formed part of the surface
of the Funafuti specimen, have also been treated by von Koch's method, but as the
soft tissues appear to be absent here, they are of value only as showing the structure
of the hard parts. The microscopic structure of the skeleton precisely resembles that
of the Lifu specimens described below.
Skeleton. Microscopic characters. The fully formed skeleton is built up of a solid
mass of polyhedral elements whose surfaces are united together to the complete ex-
clusion of the soft parts. The elements vary considerably in size, 40/it being a frequent
diameter. An exceptionally large one measures 150/i in its larger diameter. Minute
rounded granules are often present at the centre, but the remainder of the element
consists of radially disposed crystalline fibres which terminate peripherally in contact
with the fibres of adjacent elements (Figs. 3, 4, 8, and 12).
Mineralogical properties. My friend Mr A. Hutchinson, Fellow of Pembroke College,
has been so good as to examine a section and also some fragments of the skeleton
from the mineralogical point of view, and furnishes me with the following reports on it.
"Under the microscope the section exhibits well-marked .spherulitic structure and,
when examined between crossed Nicols in parallel light, each spherule is seen to be
occupied by a black cross and a system of concentric coloured rings, forming an optic
picture similar to that observed when a plate of a uniaxal crystal, cut perpendicularly
to the optic axis, is viewed between crossed Nicols in convergent light.
" Examined with a ;J-undulation plate of mica, these optic pictures behave as if
they were produced by negative uniaxal crystals.
" The double refraction of the substance is strong.
" The specific gravity, determined by suspending a small fragment of the substance in
bromoform, was found to approximate to that of Aragonite. Qualitative chemical analysis
showed that the substance is calcium carbonate. Magnesium was tested for with care,
G2— 2
464 ASTROSCLERA WILLEYANA, THE TYPE OF A NEW FAMILY OF SPONGES.
but no certain indication of the presence of that element could be detected in the small
portion at my disposal.
" All the properties described above lead to the conclusion that the substance is
composed of aragonite."
The Funafuti specimen furnished a larger fragment for analysis and determination
of the specific gravity, and of this Mr Hutchinson has made a further examination, with
the following confirmatory result : —
"A small fragment was placed in bromoform, and the liquid diluted with benzol till
the substance sank. The tube was now connected to an air-pump, and gently warmed
until the liquid boiled under diminished pressure ; the substance gave off a stream
of bubbles, and its density had apparently increased. More bromoform was now added
until the substance just remained suspended, and on taking the specific gravity of the
liquid it was found to be 284 at 15° C.
"Another portion reduced to powder gave similar results.
" The substance is therefore considerably denser than calcite, and as organic matter
is present, the specific gravity of the mineral constituent is no doubt greater than
that observed.
"The fragment was next dissolved, with the exception of an organic residue, in
hydrochloric acid, and tested ; it appeared to be pure calcium carbonate, and no trace
of magnesium could be detected."
The organic basis of the skeleton, which remains after treatment with acids, varies
remarkably in amount. In one specimen (to which the following description applies)
it is abundant, while in the two others it is very scanty. At the central regions
of the skeletal elements a highly refracting material remains, associated with a
deej^ly staining substance. These are disposed in radiating lines, and often give rise
to the appearance of a bright Maltese cross or a star with more than four rays,
whose rays of light and dark shift as the focus is altered (Fig. 10, sic.). The central
region of the skeletal element is often sharply limited by a circular boundary, the
limit of the deeply staining substance, and measures from 7 — 9;ii in diameter. At
the peripheral parts of the skeletal elements a less deeply staining and not highly
refracting substance remains, which may have a radial arrangement as in Fig. 6,
coming in contact at its edge with adjacent elements, and joining with them by a
well-marked dividing line. In other cases the peripheral parts of the skeletal
elements are represented by a granular matrix, without radial arrangement, and
merging, without a boundary line, into the periphery of neighbouring elements
(Fig. 10, sk.). I am unable to account for the diiference between individuals in the
amount of the skeletal matrix, or for that between skeletal elements in their structure.
It is to be observed that the round central regions of the skeletal elements are of about
the same size as the smallest spherules, which are to be found in the jelly. The
peripheral region of the element (distinguished by its low refraction and feeble staining
power of its matrix) is that which is laid down by concentric additions in the course
of growth. In sections of the undecalcified skeleton the two regions are undistinguish-
able. The organic basis of the skeleton takes a blue stain with picronignisin, in
ASTROSCLERA W'lLLEYANA, THE TYPE OF A NEW FAMILY OF SPONGES. 465
which it agrees with that of sponge spicules, and differs from that of Alcyonarian
spicules'.
The scattered linear spicules (spic.) represented in Fig. 8 occur both in the soft
parts and embedded in the skeleton. I believe them to be of foreign origin.
Origin of the skeletal elements. The gelatinous layer which invests the upper surface
is crowded with young growing skeletal elements, the small ones free and spherical, the
larger packed together like hailstones, and assuming the polyhedral form (Figs. 5, 7,
8, and 11).
The spherules take their origin in single cells of the jelly, near the upper surfiice
(Fig. 5, a — d). In the early stages of growth the nucleated granular cell body is seen
as a thin investing layer surrounding the spherule, which is from the first composed
of radiately arranged crystalline fibres. The smallest spherules that I have recognised
are Ifi in diameter, and I am inclined to think, from this fact and from the appearance
of the organic basis described above, that they are of this size when they are first
formed. As the spherule increases in size it takes up its position as an element of
the fixed skeleton, and in the course of their growth the angular spaces between
adjacent skeletal elements are completely filled in, to the exclusion of the soft parts.
The elements thus lose their spherical shape and become polyhedral. The external
surface of a spherule, in contact with the layer of soft tissue, is often beset with
radiating points, and resembles a portion of a spheraster of a siliceous sponge (Fig. o,
/' and (j). I find no trace of a skeletogenous layer of cells covering the fully formed
skeleton ; it appears probable that the cells in which the spherules take their origin,
remain in relation with them till their growth is finished, but I have only seen them
in the early stages.
The arrangement of the skeletal canals. Figures B and C are views of an approxi-
mately median section through one of the Lifu specimens. The canal system consists of
intimately anastomosing channels, the greater number of which have a direction
upwards and slightly outwards, opening on the ujiper surface and connected with one
another by abundant transverse and obliquely running communications.
In Figure A 5 a section of the interior of the Funafuti specimen is seen. Near
the centre of the cut surface a group of large canals, having a diameter of from 60
to 80/x., is cut across, whose arrangement corresponds with that of the large pores seen
at the surface of this specimen. The canals run close to one another and their general
direction is upwards and outwards, diverging from the axis of the fungiform ' original
growth.' In addition to those forming the central groups there are others running
approximately parallel with them in planes which are set radially to the axis of the
group. Moreover, the section lays open at least one canal which is approximately
transverse to this axis, and whose course thus corresponds with those radially directed
grooves, the initial stages of canals, seen in the Lifu specimen represented in Fig. 1, a.
In the rest of the cut surface smaller canals are exposed.
It appears probable that the large canals carry the ultimate trunks of the efferent
system of the sponge. I have not succeeded in recognising systems of large canals in
' For this fact, as well as for his assisitance in staining some sections for me with this reagent, I am
indebted to Mr G. C. Bourne, Fellow of New College, Oxford.
466 ASTKOSCLERA WILLEYANA, THE TYPE OF A NEW FAMILY OF SPONGES.
the sections of the Lifu specimens which I have examined, though the presence of
such a system is indicated on the surfece of the remaining specimen (Fig. 1, a). It
may be that a well-marked system of efferent skeletal canals is not developed until the
organism attains a certain size, and it seems clear, from the Funafuti specimen, that
the systems are multiplied, perhaps by branching of those already existing, as the
size is further increased.
In the basal region the canals are less numerous than at the upper surface, a
Fig. B. Photogii.iph of part of an appboxi-mately median longitudinal section, bt reflected light.
The skeleton (white) is seeu to be made up of separate elements. The skeletal canals (dark) are occupied
hy the (stained) soft tissue.
result produced, as sections through this region show, by the closure of the canals by the
growth of the skeletal elements which border them (Fig. C, act). The elements increase
on the surface turned towards the canal till they come in contact with those of the
opposite side. Some canals in the base remain jjermanently open, and from these the
soft tissues are apparently withdrawn towards the growing surface.
It may be well to mention here, what will be evident when the soft tissues are
considered, that though the large canals forming the systems above described are pro-
bably mainly efferent channels, the remaining skeletal canals, and by far the majority,
contain both afferent and efferent trunks of the water-carrying canals of the sponge.
Mode of Growth. Fig. C represents a section through the specimen from which
Fig. 2 is drawn, whose axis has curved in the course of growth. The section shows
that the layer of growing tissue is not limited to the upper surface, but has, as
it were, overflowed for a short distance down that side towards which the growth
ASTROSCLERA WILLEYANA, THE TYPE OF A NEW FAMILY OF SPONGES. 467
'^ssm^'-''
^M'
Fig. C. An APPB0XI^UTELY median longitudinal section thkough the specimen kepbesented in Fio. 2.
The skeletal canals in the upper part are seen to contain soft tissues, while those at the base are
empty. (The canals to the left, marked by oblique shading, contained broken fragments produced by sawing,
so that the nature of their contents could not be recognised.) a, a. Canals which have been filled in by the
growth of the skeletal elements bordering them. b. Region of the skeleton formed by overlapping layers of
skeletal elements added on the outer surface.
468 ASTROSCLERA WILLEYANA, THE TYPE OF A NEW FAMILY OF SPONGES.
has tended. An examination of the base of this side {b) shows that this process has
occurred many times in the earlier stages. It appears that the upper (perforated)
surface of this specimen has shifted in the course of growth, the canals becoming
closed on one side while new ones have appeared on the other. The layer of cortical
(unperforated) skeleton which may be seen in the Funafuti specimen (Fig. A, 1) to
extend from the bases of the later formed bosses over the surface of the original
growth has probably a similar origin.
The specific relationship of the specimens from two localities. As the soft tissues
proper to the organism are absent from the Funafuti specimen the question of its
specific relationship to those from Lifu may here be considered. Do they belong to
the same species, of which those from Lifu represent the younger and that from Funa-
futi a more advanced stage, or are they specifically distinct ? The scanty supply of
material prevents a definite answer being given, but the following points may be urged
in favour of the former view :
The characters of the skeletal elements are, I believe, identical.
Though the shapes of the organisms from the two localities are very different, there
is to be seen in the Lifu specimens a tendency to expand as they grow, which if carried
out might well lead to the formation of a structure like that which is called above
the original gi'owth of the Funafuti specimen. A similar change of shape occurs in
the coral Fungia, which in the young state is cylindrical and later expands into
a disc.
The difierentiation of the efferent (?) canal system appears to be more complete
in the Funafuti specimen, but this is a character which might, as above suggested,
become more marked in proportion as the extent of the system to be drained increased,
and one of the Lifu specimens (Figs. I and 1, a) shows a distinct indication of it.
The fact that the Lifu specimens (as stated below) contain eggs is an indication
that they have attained sexual maturity, but not that they are at the limit of their
growth.
It appears that there is at present no sufficient ground for regarding tlie forms
from the two localities as specifically distinct.
The Soft Tissues. The gelatinous layer above described as investing the ridges
of the skeleton, lines the openings of the canal system and extends as a sheet,
thinner in the centre, over the end of each canal (Fig. 8). A round pore is
frequently present opposite the centre of a canal (Figs. 7 and 11), but in many cases
the membrane appears not to be perforated at this point. The spaces at the mouths
of the canals are however in communication with one another, by lateral channels
through the jelly, beneath the surface membrane.
The soft tissue can be traced for a short distance down the sides as a thin
layer, investing the outer surface of the animal.
Besides the cells in which the spherules are formed, there are branched amoeboid
cells .sparsely scattered through the jelly (Figs. 18 and 19). The jelly, as well as the
organic basis of the skeleton, takes a faint blue stain with picronigrosin.
ASTROSOLERA WILLEYANA, THE TYPE OF A NEW FAMILY OF SPONGES. 469
The soft tissues of the interior of the sponge are contained in the skeletal canals
and penetrated by the water-bearing canals which open by the pores at the surface.
From the main canals small ones are given oft" which ramify in the layer of soft tissue
in contact with the skeleton.
As the canals are followed downwards into the interior, the cellular elements in
their walls, at first scattered, become more and more abundant, and the jelly less
conspicuous. In some parts of this intermediate region large coarsely granular cells
are abundant in the jelly. At a short distance from the surface the soft tissue
assumes the characters which are maintained throughout the interior (Fig. 12).
The appearance of a section through it (Figs. 14 and 15 — 17) suggests that it
is largely made up of cells united into a reticulum, with vacuolar spaces of various
sizes forming the meshes. The jelly, which is so conspicuous in the surface layers,
appears to be scanty or altogether absent here. Besides the smaller, branched proto-
plasmic masses, with small nuclei (1".5 — 2fi in diameter) which make up the greater
part of the reticulum, there are larger and more circumscribed cells with larger nuclei
(2 — 3/x). Scattered through the reticulum, and with their walls apparently formed by
portions of it, are ciliated chambers and the ramifying branches of the canal sj-stem.
Ciliated chambers (Figs. 15 — 17). The.se are round or oval chambers of minute
but fairly uniform size, the larger measuring 18 by llfi, the smaller 10 by 8/i. (The
sections of smaller diameter are doubtless, in many cases, transverse to the long axis of
larger chambers.) Their walls appear to be formed by cells which send out processes
laterally, and these, joining with one another, bound the chamber; other processes
extending away from the chamber are continuous with other cells of the reticulum,
while a thii'd set of processes project into the cavity of the chamber, and each,
tapering gradually from its base, forms a flagellum which may extend across to the
other side of the chamber. There is no indication of a collar or of the abrupt truncated
termination of the cell-body at the base of the flagellum, which are usually seen in
choanocytes. A well-marked nucleus is situated at the base of the flagellar process, at
its junction with the body of the cell. The flagella project from about half the inner
surface of the chamber, some four or five commonly appearing in section, and their
tips thus converge and are often seen to have become entangled. The remainder of
the inner wall of the chamber is smooth and at some point in it the cavity opens
into a branch of the canal system by a narrow passage, about 4/i in diameter (Fig. 17).
This is only rarely seen as a distinct passage, owing to its narrow lumen.
Considering how readily the collar cells of sponges lose their characters unless
special measures are taken to preserve them, it cannot be assumed that the condition
above described of the cells lining the ciliated chambers is precisely that of the
living state. It is perhaps possible that they had in life the usual characters of
collar cells, but if so. it is curious that while the flagella are preserved they should have
lost not only their collars, but also the truncated shape of the ends turned towards
the centre of the chamber. And, in view of the unique character of the skeleton of
Astrosclera it is quite possible that the cells of its ciliated chambers were also of a
peculiar type. We cannot, with the present material, arrive at certainty on the pouit.
w. IV. 63
470 ASTROSCLERA WILLEYAXA, THE TYPE OF A NEW FAMILY OF SPONGES.
All I can say is that in the material, preserved as it has been, the condition appears
to be as I have described.
Canal System. The large trunks of the canal system and the small branches
which ramify in the layer of tissue containing ciliated chambers have nuclei scattered
uniformly in their walls (Fig. 12, canal to right, and Fig. 14, c), but I have not suc-
ceeded in recomising cell limits about them.
Most of the skeletal canals are lined by tissue containing ciliated chambers in
abundance, though in some rare instances, as in the left-hand canal represented in
Fig. 12, the lining contains no ciliated chambers. It may be that these are efferent
passages, but I have not succeeded in recognising in the Lifu specimens, (which alone
contain the soft tissues,) a clearly differentiated system of large canals corresponding to
those seen in the Funafuti specimen.
The existence of ciliated chambers implies the presence of systems of afferent and
efferent canals ; and as a matter of fact we find that the tissue containing the ciliated
chambers is traversed by trunks which give off small branches into the tissue. Of the
small branches seen in the tissue some may well be tributaries of the efferent system
of canals.
It appears that in this young stage at least, the main trunks of the efferent
system are in many cases also surrounded by tissue containing ciliated chambers,
together with efferent tributaries and the ultimate branches of the afferent sj-stem, so
that in sections of the soft parts the two systems of canals are not recognisable by
their anatomical characters.
Reproduction. Each of the tliree specimens examined contained large eggs or
embryos (Figs. S, 9 and 10). They are found, solitary or two together, near the orifice
of one of the larger canals, separated from the skeletal wall by a thin layer of soft
tissue. I have not been able to recognise eggs in a young stage. An advanced ovum
in one specimen (Fig. 9) measures O'l mm. in length, and has a thick-walled nucleus
25/i in diameter, and a well-marked germinal spot. An embryo in the same specimen
(Fig. 10) is rather larger than the egg. At the surface there is a superficial layer of
nuclei, and the protoplasm about them is disposed in columns perpendicular to it.
Internally the columns are merged in the granular protoplasm, which occupies the
interior of the embryo, obscurely divided up into irregular masses, but I have not
succeeded in detecting nuclei in them. None of the embryos have a segmentation
cavity. It appears that the development leads to the formation of a larva of a
parenchymula type, rather than an ainphihlastula.
I have not been able to recognise any stage in the formation of spermatozoa.
State of Preservation. The three specimens whose soft parts have been examined
do not show the structure equally well. The above description is given from the
Lifu specimen which was prepared by von Koch's method, and of which part was
cut into sections 5/i in thickness. In another specimen the reticular character
of the soft tissue is clearly seen, but the fiagella of the ciliated chambers are
obscure. This is partly due to the fact that the sections are in .this case considerably
ASTROSCLERA WILLEYANA, THE TYPE OF A NEW FAMILY OF SPONGES. 471
thicker than the diameter of the ciliated chambers, so that the latter caiiuot be
seen actually in section. There is however another fact about this specimen which
may possibly explain the obscurity in the structure of its tissues. The canal system
contains in many parts hosts of minute deeply staining rod-like bodies (1 — ofi in
length), looking very like bacteria (Fig. 20, h.). They occur in scattered gi-oups or
apparently embedded in some clear homogeneous substance, in such masses, that they
appear to distend the canals. Three possibilities as to their nature have occurred
to me, namely, that they are bacteria, spermatozoa, or the remains of food. They
are not symbiotic bacteria, for they are absent from the two other specimens. The
cri.sp, evidently growing surface of this specimen, and the excellent preservation of
the eggs and embryos (those represented in Figs. 9 and 10 were contained in this
specimen), seems to put the view that the animal was undergoing decay, out of the
question. I do not think that the bodies are spermatozoa, for I have found no trace
of sperm morulae or any of the earlier stages of development of .spermatozoa. If they
are the remains of food it is remarkable that they should be found only in one
specimen, and I am moreover entirely at a loss to form a conjecture as to the
nature of the food which would leave such remains.
Affinities.
That the animal under consideration belongs to the Porifera seems clear from
the presence in the soft tissues of chambers provided with flagella and communicat-
ing with a system of canals which ultimately open to the exterior by pores. The
soft tissues are moreover supported by a skeleton composed of elements secreted by
cells scattered through the jelly.
On the other hand that it is not a Coelenterate is shown by the absence of
polyps, mesenteries and thread-cells.
There is at first sight some resemblance between a section of the skeleton of
Astrosclera and sections in certain planes of the skeleton of the Madreporarian Corals.
In these the fibres of the skeleton are disposed in fascicles and diverge from one
another about an axis which occupies the centre of the fascicle. A section which
cuts the fascicles transversely thus presents the appearance of fibres radially disposed
about a number of centres. Such a section is shown in Fig. 10 and other figures
of Miss Ogilvie's memoir " On the Microscopic and Systematic Study of the Madre-
porarian Types of Corals'." But, as Miss Ogilvie clearly shows, these systems of
radiating fibres are made up of lamellae successively deposited on the surface of the
skeleton, so that any particular fibre of a fascicle traverses many lamellae in its
course. We know from the work of von Koch, von Heider and others that the
skeleton is formed by a layer of cells, the calycoblasts, in contact with its surface.
It will be readily seen that the resemblance between these radiating systems of
fibres, and those formed within single cells of Astrosclera, can only be superficial.
' Philosophical Transactions, vol. 187, B (18'JG).
63—2
472 ASTROSCLERA WILLEYANA, THE TYPE OF A NEW FAMILY OF SPONGES.
In von Koch's account of the development of the skeleton in Asteroides caly-
cularis^ its elements at their first appearance are described as spheroidal bodies, which
become polyhedral by apposition and possess a radiate structure (PL 20, Fig. 7). They
have the appearance of being partially divided into two pieces. Von Koch regarded
these bodies, as well as the later formed skeleton, as excretions, formed by the calycoblast
{epiblast) cells of the coral.
The precise relation of the calycoblasts of corals to the skeleton which they form is
however still uncertain. The question is whether the skeleton is, as von Koch and other
authors have supposed, secreted by the layer of cells in contact with it, or built up of
cells of this layer which themselves become calcified. Miss Ogilvie urges strong reasons
in favour of the latter view.
In either case it is not clear what the relation of the spherules described by
von Koch to the fully-formed skeleton may be, for in this such spheroidal elements do
not appear to be present.
I mention these spherules here because they are skeletal elements which corre-
spond in composition (aragonite), and to more extent in structure, with those of
Astrosclera, but not as evidence of any close affinity of the latter with the Coelen-
terates.
But, admitting that Astrosclera is a sponge, there are many features which
separate it from the living members of this group. Among these may be mentioned : —
(a) The shape of the skeletal elements. They are polyhedra which begin in a
spherical and may pass through a spheraster stage.
(6) Their union to form a rigid skeleton, excluding the soft parts. In Petrostroma
Doderlein, the representative of the Lithonina'-, the supporting skeleton is formed of
fused spicules, but these are modified quadriradiates. We are at present without
information on the soft tissues of this form.
(c) The mode of growth, by the addition of new skeletal elements at the upper
surface, and without interstitial growth. In this feature also it probably agrees with
Petrostroma.
(d) The limitation of the pores to the upper surface.
Tentorium (Vosmaer), a siliceous sponge classed with the Polymastidae, presents a
similar limitation.
(e) The absence of a central atrial space.
(/) The small size (18 by 11/i) of the ciliated chambers. The smallest size given
by Haeckel for the ciliated chambers of the Leucones (in which gi-oup they are smaDer
than those of other Calcarea) is 60 by 40/x. Among the Non-calcarea, 42/a is the
smallest diameter that I have found given for the ciliated chambers.
1 "Ueber d. Entwickelung d. Kalkskeletes von Asteroides calycidaris u. dessen morphologische Bedeutimg."
Mitth. aiis d. Zool. Stat, zu Neapel, Vol. 3, 1882, p. 284.
' Doderlein, L., "Ueber die Lithonina, eine neue Gruppe von Kalkschwammen. " Zool. Jahrbiicher, Syst.
Abth. X. (1898), p. 1.5.
ASTROSCLERA WILLEYANA, THE TYPE OF A NEW FAMILY OF SPONGES. 473
(g) To these points of difference is perhaps to be added the character of the
flagellated cells lining the ciliated chambers. They appear, though the point needs
confirmation from specially preserved specimens, not to be collar cells of the ordinary
type, but more or less amoeboid and without a collar, and with one of their processes
gradually tapering into the flagellum.
From the Calcarea Astrosclei'ci also differs in the following features : —
(a) The flagellate cells are limited to about half the interior of the ciliated
chambers.
(b) There appears to be a long and complex canal system both on the afferent
and efferent sides of the ciliated chambers.
(c) The mineral constituent of the skeleton is arar/onite, not calcite.
Comparison of Astrosclera 7vith the Pharetrones.
The Pharetrones are calcareous sponges which are found in beds ranging from
the Carboniferous to the Cretaceous formations. In the arrangement of their skeleton
they differ widely from living sponges except Petrostronia the type of the Lithonina.
Dr Hinde has recently described' some sponges from the Eocene beds of Australia
which appear to be intermediate forms connecting the Lithonina and the Pharetrones.
The resemblance in general characters between Astrosclera and some of the
Plmretrones appears to me so striking as to deserve consideration, although I have
to conclude that the differences in the minute structure of the skeleton forbid their
being considered as really allied.
The feature characteristic of many members of this group (though authorities are
by no means agreed on its essential cliaracters or limits) is that the spicules are united
in close apposition to form trabeculae or ' fibres,' as they are technically called. Dr
G. J. Hinde writes, " in no existing sponge is there the same disposition of the spicules
of the interior to form anastomosing solid fibres as in this and other allied fossil
forms ; and this character forms the distinctive feature of the Pharetrones, sharply
marking them off from the families of existing Calcisponges."'
The centre of the sponge is often occupied by a space, the gastral cavity, with
a wide opening above, and penetrating more or less deeply into the interior. The
system of anastomosing canals which lies between the trabeculae of the skeleton
communicates on the one hand by fine pores with the exterior, and on the other,
either by irregularly anastomosing channels or by definite tributary trunks, with the
gastral cavity. In some cases the gastral cavity may be so shallow as almost or
altogether to cease to exist, and in that case a group of large tributary trunks opens
directly to the exterior or at the bottom of a shallow depression. A cortical layer, which
has been thought to be imperforate, often clothes the sides of those members of the
group which are pedicellate or cylindrical, and if this is so the pores opening to the
1 G. J. Hinde, " Calcisponges from the Eocene of Victoria {.Australia)." Quart. Joiirn. Geol. Soc, Vol.
LTi. 1900, p. 50.
» "Notes on Fossil Calcispongiae, " .Inn. and .Mag. of Nat. Hist. Ser. 5, Vol. x. (1882), p. 190. This
was written before the discovery of the Lithonina.
474 ASTROSCLERA WILLEYANA, THE TYPE OF A NEW FAMILY OF "SPONGES.
exterior are limited to the upper part of the sponge. In some cases radiating grooves
are present on the upper surface, converging towards the mouth of the gastral cavity,
or towards the group of large pores which represents it, the initial stage of radiating
tributary canals of the interior.
In the genus Stellispongia the gastral cavity may, as above noted, be absent
altogether, and a single group of large pores and radiating grooves may occupy the
centre of the upper surface of the sponge. In other cases, and even as it appears
within the limits uf the same species, the number of these groups of pores may be
multiplied.
Among a number of specimens of Stellispongia vanabilis which I obtained from
Triassic deposits in the neighbourhood of St Cassian, in the Tyrol, and which have
been kindly named for me b}' Dr Hinde, there are specimens, referred to the variety
c of that species, expanding rapidly from a short pedicellate base, clothed on the
outer surface with a wrinkled cortical layer, and presenting in the middle of the upper
surface a shallow depression whose sides are marked by radiating gi-ooves, the first stage,
as we have seen of the tributary canals of the interior. In another example of the
species the rounded upper surface presents numbers of these systems of large pores
and radiating grooves, in some cases in depressions, in others flush with the general
surface.
The resemblances between the surface of the latter specimen and that of the bosses
of the Funafuti specimen of Astrosclera, and between the general habit of growth of the
former specimen and the earlier formed part of the Funafuti specimen are certainly
very remarkable.
On turning our attention, however, to the elements of which the skeleton is
composed we find a marked difference. In those Pliaretrones in which their characters
have been determined they are tri-radiate or quadri-radiate spicules, often haviuo- the
rays bent or reduced to allow of their being packed together to form the solid
trabeculae, but still distinctly referable to these types. They are well seen in the
Warminster specimens described and figured by Dr Hinde', who succeeded in obtaining
isolated spicules. The characters of the skeletal elements are however often found to
have been obliterated to a greater or less extent in the process of fossilization and
all traces even of the existence of spicules may be lost.
In sections of the specimens of Stellispongia above mentioned the skeleton is seen
to be composed of bodies round in transverse section, but elongated in longitudinal
with a length often eight times the breadth, and with no definite structure discernible
in their interior. The appearance agi-ees with that figured by Steinmann for this
genus-. The outlines of the elements which make up the trabeculae are blurred, owing
apparently to commencing recrystallization, so that it is not easy to decide on their
original shape. Rauff^ asserts that tri-radiate spicules are to be recognised in some
specimens of Stellispungia from the Trias, and in my sections the elements often present
' "Notes on Fossil Calcispongiae," loc. cit.
' Cp. Steinmann's figure, " Pharetronen-Studien," Neiies Jahrbiich j. ilineralogie, &c. 1882, Bd. 2, PI. ix.
Figure 2.
^ Palaeos-pongiologie, Th. 1. PalaeontO(iriqihica, Vol. xl. p. 99.
ASTROSCLERA WILLEYANA, THE TYPE OF A NEW FAMILY OF SPONGES, 475
an augular beod in their course, which is iiuite consistent with this statement. It is
probable then that the skeleton of Stellispongia is of the same spicular type as that of
other Pharetrones.
It cannot, therefore, be claimed that there is any resemblance between these spicules
and the polyhedral elements of Astrosclera.
We are confronted then with the fects (1) that there have existed a group of
sponges, the Pharetrones, with the above-mentioned characteristic features of their
anatomy and a skeleton composed of spicular elements, and (2) that Astrosclera
exists at the present da}' approaching some members of the Pharetrones closely in the
general features of its anatomy, but with a skeleton of polyhedral elements.
Xow the characters of the elements of the skeleton are those which are mainly
relied on in the classification of sponges, whether of the large gi-oups, as the names
Hexactinellida, Tetractinellida, Monaxonida imply, or their subdivisions.
As regards form, the difference between the skeletal elements of Astrosclera,
whether in their earl}- spherical or spheraster-like state, or in their later shape of
polyhedra, and the tri- or quadri-radiate spicules of the Pharetrones is as great as can
be foimd among the spicules of sponges.
As regards develojnnent, the best clue we have to the way in which the spicules
of the Pharetrones were formed is furnished by Minchin'.s observations on the growth
of tri-radiate and quadri-radiate spicules in Calcareous sponges at the present day.
Minchin finds, and I have myself had the opportunity of examining his beautiful
preparations, that a tri-radiate spicule is laid down by a group of six cells, derived
originally from three, each of which give rise by divi-sion to a pair which preside over
the formation of one ray of the spicule. In Astrosclera, as we have seen, the
spherules of the skeleton are contained at their first appearance in a single cell.
Finally, as regards composition, the carbonate of lime is in Astrosclera in the form
of aragonite, while, from the analogy of the spicules of the Calearea of the jiresent
day, we may suppose that those of the Pharetrones were of calcite.
To judge then by the rules which have become established in classifying sponges,
we must conclude that Astrosclera is a representative of a distinct family, and not
closely allied to the Pharetrones. The resemblances between the two groups may on
this view be ascribed to the similarity of the conditions under which they have growm.
On the whole, I believe that this is the correct conclusion, though the resemblances
are so striking that I can hardly persuade myself to admit that they can be only
adaptive.
To fortify myself in this conclusion I have to reflect on such cases of similarity
in general habit between widely separated groups as that shown by some members of
the Euphorhiaceae and the Cactaceae, ordei-s which no botanist would admit to be allied
to one another.
Apart from the nature of the elements constituting the trabeculac I do not
know what characters could be given to distinguish Astrosclera from some of the
Pharetrones such as Stellispongia, and it appears very possible that among the forms
included in this group whose microscopic characters have not been ascertained, there
may be some with polyhedral skeletal elements, truly allied to it.
476 ASTROSCLERA WILLEYANA, THE TYPE OF A XEW FAMILY OF SPONGES.
On the Spherulitic Structure seen in Pharetrones from St Cassian and elseiuhere.
In the triassic deposits of St Cassian and others occurring near Schhiderbaeh and
referred to the same horizon, representatives of the Pharetrones occur in which the
carbonate of lime trabeculae of the skeleton show no trace of spicules, but, instead,
the\' are made up of more or less polyhedral systems of radiating lines which at their
periphery are in contact with adjacent systems either by a sharp or a confused
boundary'. They thus remarkably resemble the trabeculae of the skeleton of Astrosdera.
The view of this structure held by Zittel, and by most palaeontologists who
have considered the matter, is that it is secondary, being due to a recrystallization
of the lime. I have carefully examined sections of specimens which I obtained from
these localities, in several of which the sphaerulitic structure is well marked, and I
am prepared to accept this conclusion as correct.
The evidence bearing on the matter may be arranged under separate heads.
a. If the spherulitic structure of the fossils were primary we might expect that
those in which it is present would form a group differing in other respects from the
Pharetrones with a spicular skeleton. The opposite is the case.
Among the genera included in the group there are simple forms, of which Stelli-
spongia and Corynella may stand as examples, and remarkable segmented forms con-
sisting of hollow segments superposed one upon another, so that the roof of one forms
the floor of its successor. These have been divided by Steinmann as a separate group,
the Sphinctozoa, and Zittel regards them as so distinct that he removes them from the
Pharetrones, and classes them in a separate order. The precise relationship of the two
groups does not materially affect our present enquir}-.
The genus Verticillites is one of these segmented forms. In a representative of
it occurring in the Greensand beds of Warminster (F. D'Orbignyi, Hinde) the com-
position of its walls of tri- and quadri-radiate spicules has been demonstrated as clearly
as in any recent sponge. (Cp. Hinde, Ann. and Mag. ^. H. 5, x. PI x. Fig. 3 &c.)
On the other hand, the genera Thaumastocoelia and Enoplocoelia of Steinmann are
examples of the segmented forms from the triassic strata of St Cassian, and the
specimens which have been examined present, wholly or in part, a spherulitic structure
of the walls. (The structure was regarded by Steinmann as primary in the fcjrmer
genus, but as secondaiy in the latter.)
Again, the genus Corynella is one of the unsegmented forms and possesses distinct
characters'-.
' The structure is figured by Zittel, " Studien iiber fossile Spongien, in.," Abh. d. ilath.-Phys. Classe d. kon.
bayerischen Akad. d. Wisseiischaften, Bd. xiii. Abth. 2, PI. xii. Fig. 5 {Corynella gracilis), also in his GrundzUge
der Palaeontologie, p. 59, Fig. 88.
- According to Zittel's description (" Studien iiber fossile Spongien," p. 3-5) they are usually solitary
sponges, cylindrical or pear-shaped, thick-walled, and with a truncated or rounded upper surface. The gastral
cavity is funnel-shaped, rarely reaching to the base. Eadial grooves are often present round its mouth and
wide tributary tubes pass outwards and downwards from the gastral cavity. As they penetrate the walls
they become narrower and ultimately open by fine inhalent ostia at the surface. A thick dermal layer is
sometimes present at the base. The skeletal fibres are coarse, chiefly composed of simple spicules, but with
some large tri-radiate ones.
With regard to the 'simple' spicules Hinde points out that those of C. socialis are tri-radiates so
modified that they might very readily be mistaken for simple ones.
ASTROSCLERA WILLEYANA, THE TYPE OF A NEW FAMILY OF SPONGES. 477
In C ruffosa, from the Upper Greensand of Warminster, Hinde succeeded in obtain-
ing complete and isolated tri-radiate spicules (loc. cit., PI. XI. Fig, 2.5), and sections of
well-preserved specimens of this genus commonly show the ' fibre ' to be composed of
such spicules.
In Zittel's figure of a specimen of C. gracilis, Mlinstr. from St Cassian, alluded
to above, no trace of sjjicules is to be seen, and the ' fibre ' has instead a well-marked
spherulitic structure.
Now on the view of the pi-imary nature of the bodies with radiating lines, making
up the spherulitic structure, we are driven to suppose that among the sponges which
have been regarded as Pharetrones there are two groups, one with spicular skeletal
elements, the other with polyhedral skeletal elements and that both have developed iuto
simple and segmented forms. It was shown in the last section that in its larger
characters Astrosdeni does remarkably resemble such unsegmented forms of the
Pharetrones as Stellispongia. But that there should also be segmented forms with
polyhedral skeletal elements resembling the segmented Pharetrones would indeed be an
astonishing instance of parallel development.
b. Many intermediate conditions are met with between a ' fibre ' composed of
spicular elements and one exhibiting the spherulitic structui-e. Thus the fossils
belonging to the genus Gorynella have, as we have seen, in many cases, a fibre com-
posed of spicules. In a section of a specimen of C. forarninosa from the Lower Green-
sand of Farringdon\ while the spicular elements are clearly seen in the centres of the
fibres, their peripheral parts present a fine crystalline striatiou perpendicular to the
surface, and there is no sign of spicules in these regions.
Again, though the spherulitic structure is so clearly marked in the specimen of
Gorynella gracilis Mlinstr. above alluded to, in one of my specimens (from Schluderbach)
referred by Dr Hinde to this species, the central parts of the trabeculae are made up
of elongated bodies which may well be the remains of spicules, though here too there
is at their periphery a crystalline striation more or less perpendicular to the surface,
though the lines are in part disposed in groups with a fan-like radiating arrangement.
Similarly, in Pharetrospongia Strahani Sollas, the ' fibre ' is often seen to contain
spicules lying, as in a quiver, in a direction parallel to the .surface, but in one of
Dr Hinde's sections of a specimen of this species, no trace of spicules is to be seen,
and instead, the 'fibre' presents lines .sometimes disposed perpendicularly to the surface,
sometimes extending in fan-like and frond-like systems, with definite boundaries between
thcni, recalling the patterns on a window-pane in frosty weather.
If the spherulites with fibres radiating in all directions are regarded as primary
elements of the skeleton, what, it may be asked, is to be made of the imperfect
systems, and of these borders of fibres set nearly perpendicularly to the surface of
the trabeculae, which are of much more frequent occurrence ?
c. Although, in its more marked form the spherulitic structure does remark-
ably resemble that seen in Astrusclera there is, I believe, this difl:erence ; that whereas
the radiating lines in the latter run straight from the centre to the circumference of
> Kiudly lent to me by Di- Hinde.
vv. IV. 6't
478 ASTROSCLEBA WILLEYANA, THE TYPE OF A NEW FAMILY OF SPONGES.
the skeletal elements, in the former the diverging lines are frequently grouped in
frond-like systems, whose axes radiate from the common centre.
d. Even in the same specimen the spherulitic structure may be found in one
part of a specimen and the spicular structure in another. Thus Zittel speaking
of the spherulitic structure, says : — " Auch dieser Erhaltungszustand kommt zuweilen
an Exemplaren vor, bei denen sich einzelne Fasern noch deutlich als aus Nadelu
bestehend erweisen." (Studien ilher fossile Spongien, lir. p. 14.)
e. In a specimen of CoryneUa tetragona tigured by Zittel (PI. XII. Fig. 2) the
trabeculae are composed of apparently simple spicules embedded in a matrix, but the
spherulitic structure is also present, printed as it were over the other, and with its
radiating fibres pursuing their course regardless of the presence of the spicules.
/. In some of my St Cassian specimens the spherulitic structure Is limited to
the trabeculae, but in many cases it extends out from them into the calcite mosaic
which fills in the original spaces of the sponge, and in some cases spherulites occur
distinct altogether from the trabeculae and isolated in the calcite filling the spaces.
There is one argument which has been brought forward in favour of the primary
nature of the systems making up the spherulitic structure, which deserves further
considei'ation. Steinmann describes in the wall of Thauinastocoelia which has this
.structure, minute ramifying passages which he believes to be Thallophyte borings, and
appeals to their presence in support of the view that the parts which they penetrate
had their present structure at the time that the borings were made\ If this con-
conclusion were justified, I do not see how the argument could be met. Rauff-
doubts the interpretation which Steinmann gives of these markings, and expresses his
belief that they indicate the remains of spicules. What their real nature may be in
this particular case I am, of course, unable to say, but in one of my own specimens of
a Pharetronid fi-om Schluderbach, the spherulitic structure is very clearly marked, and
the systems of radiating lines are traversed by ramifying markings resembling those
figured b)' Steinmann, and certainly not the remains of spicules. They precisely resemble
the borings which are found in shells and corals at the present day, made by an
organism sometimes referred to the genus Achlya^. The conclusion, however, that the
presence of these borings is evidence that the skeleton which they penetrate has retained
the same structure that it had when the borings were made seems to me to be quite
uncertain.
On the view that the spherulitic structure is the result of the recrystallization
of the lime, replacing the original spicular structure, it is not to be supposed that
the S3'stems of crystalline fibres have mechanically pushed their way from the centres
through the trabeculae of the skeleton to the destruction of any delicate bodies that
1 Steinmann G. " Pharetroneu-Studien." Nnuex Jahrbuch j'i'ir Mineralogie, &c. Jalirg. 1882, Bd. ii. p. 139.
The passage alluded to is on p. 186, "Es liegt auf der Hand..."
^ Palaeospongiologie {Palaeontographica, Bd. XL.), p. 100.
^ Achlya is one of the Saprolegnieae. The grounds for this view of the relationship of the organism seem
insecure.
ASTROSCLERA WILLEYAXA, THE TYPE OF A NEW FAMILY OF SPONGES. 479
might be present, but that they result from slow molecular changes set up in the
trabeculae in continuity with the parts already changed.
The whole question of the cause of the change and the way in which it has
operated is, of course, very obscure, but I have had the opportunity of seeing what
appears to be an instance of a somewhat similar process in a specimen of fossil wood
in the possession of my friend Mr Seward, and ligured by him in his work on Fossil
Plants (Fig. 14, B, p. 81). The siliceous matrix of the specimen has at certain points
taken on a definite structure, showing concentric alternating rings of light and dark.
In longitudinal sections the structure of the woody fibre is obscurely seen in the regions
outside the systems of concentric lines, but within them, precisely where the change in
the matrix has occurred, the structure is preserved in exquisite perfection, so that the
minute transverse striations on the walls of the vessels are clearly seen.
There seems therefore to be good reason for believing that in the case of the
St Cassian sponges the spherulitic structure is the result of recrystallization and
masks a primary spicular structure. The possibility must however be borne in mind
of the occurrence of Astrosclera or allied forms in a fossil state. If the fossils retained
theii- original structure they would differ from the St Cassian sponges which I have
seen, in the absence of traces of a spicular skeleton, and also, I believe, in the strictly
radial character of the fibres comj^osing their skeletal elements, and in the uniformity
and completeness of the radiating systems thus formed.
From one point of view the resemblance between the radiate structure of the
skeletal elements of Astrosclera, and that of the spherulitic systems of the fossils
is interesting ; namely that it is an instance of the formation in living protoplasm
of a skeletal structure composed of carbonate of lime in a form almost identical
with that which this mineral may take in crystallizing outside a living body, and
thus in inorganic surroundings.
The Family Astroscleridae.
In view of the isolated position which Astrosclera holds among sponges it must
be regarded as the type of at least a separate family. It appears indeed that a
larger division will have to be established for its reception, but until specimens are
available in which the minute characters of the soft parts can be detennined with
certainty, the establishment of such a division may be deferred.
I propose to call the new Family Astroscleridae. Its characters are, at present,
those of the species which constitutes it, namely :
Sponges which are nearly cylindrical, or expand from a narrow base into a wide
disc, — differences in shape which may represent stages of growth. The upper surface
is convex.
The sides and the under surface of the disc, when this is present, are imperforate
and more or less conspicuousU' marked by annular wrinkles.
The upper surface is pitted by closely set pores, the openings of the skeletal canal
system. The pores may be irregularly distributed, or larger pores may be arranged
in one or more systems of radiating lines.
64—2
480 ASTROSCLERA WILLEYANA, THE TYPE OF A NEW FAMILY OF SPONGES.
The skeleton is rigid and built up of elements consisting of aragonite, contained
in an organic basis, and having a fibrous structure disposed in lines radiating from a
point in the centre of the element. The elements are formed in single cells and are
at first free and spherical, the}' may pass through a spheraster stage, and are ultimately
built together, becoming polyhedral by mutual apposition into a compact skeleton from
which the soft tissues are excluded.
The skeletal canals permeate the skeleton freely branching and anastomosing, and
open at the pores on the surface. In the later stages (?) of growth, large trunks of
the skeletal canal system are formed, which open at the larger f)ores at the surface.
The soft tissues form a layer at the surface, and are contained in the skeletal
canals. The ciliated chambers are minute (18 by 11 /i) and both afferent and efferent
systems of canals are long and branched. It is probable that the larger efferent
trunks run in the large skeletal canals. There is no common atrial cavity, and the
canals communicate with the e.xterior by pores in the gelatinous layer covering the upper
surface.
The larva belongs to the par enchy inula type.
Astrosclera ^villeyana occurs at depths of 3.5 to 40 fathoms at Sandal Bay, Lifu,
Loyalty Islands, and at a depth of 100 fathoms on the outer slope of Funafuti, Ellice
Islands.
The name of the genus Astrosclera is descriptive of the star-like arrangement of the
fibres in the skeletal elements, and I propose the specific name ivilleyana in commemo-
ration of Dr Willey's labours among the islands of the Western Pacific, carried on with
persistence and enthusiasm and more than once at no small personal risk, in pursuit
of the object which he had in view.
In conclusion, I wish to express my thanks to the authorities of the British Museum
for allowing me to examine the Funafuti specimen of Astrosclera ; to Dr G. J. Hinde, F.R.S.
for naming my Pkaretrones from the Tyrol, and for his kind assistance in other ways ;
and to Mr Hutchinson of Pembroke College, for his careful determination of the physical
and chemical properties of the specimens of the skeleton of Astrosclera which I placed
in his hands.
Wi'.r.KY Zoological Results.
Plate XLV
J.J.Ii&ter del.
LISTER. ASTROSGL.KRA.
WestjWswman Irth
DESCRIPTION OF PLATES XLV— XLVIII.
(Lister. Astrosclera.)
REFERENCES.
c. canal. <•'. small canal.
c. ch. ciliated chamber.
c.j. scattered cells of the jelly.
cort. cortical layer of the skeleton.
/. c. large cells of the soft tissue.
in. surface membrane.
p. pore.
sk. skeleton.
sk. e. skeletal element.
spli. spherule, young stage of skeletal element.
spic. spicule (of foreign origin).
All the Figures ichich Jolhno are draicii from specimens Jroin Lifxi.
Plate XLV.
Fifi. L Side view of a specimen S mm. in height.
Fk;. Ire. View of the upper surface of the same specimen.
Fig. '1. Side view of a specimen which has curved in the course of growth, so that
its axis comes to be parallel with the surface of attachment. Height 10 5 mm.
Fig. 2«. View of the upper surface of the same specimen.
Fig. 3. Part of a chip broken from the skeleton, x 570.
Fig. 4. Four skeletal elements, from a thin section, x 690.
Fig. 5. Skeletal elements from the upper surface in difl'erent stages of growth, x 690.
rt. One of the skeletogeuous cells of the jelly, containing a small spherule 8/j. in
diameter. Other cells of the jelly are also seen.
6 — (/. Growing spherules, invested by the skeletogenous cells, n. Nuclei of the latter.
e. A growing spherule still free in the jelly.
f. Spherules becoming adherent and with flattened adjacent surfaces.
(J. A skeletal element in the spheraster stage.
Fig. 6. Section of the organic basis of the skeleton of a decalcified .specimen. x 570.
At b a skeletal element with an unusually large centre is shown.
Fig. 7. Two of the depressions in the upper surface (represented in Figs. Ire and 2a)
seen from above and illuminated by direct sunlight, b. is seen obliquely. The surface mem-
brane is shown perforated by pores (p.) and containing spherules.
482 ASTROSCLERA WILLEYANA, THE TYPE OF A XEW' FAMILY OF SPOXGES.
Plate XL VI.
Fig. 8. Part of a vertical section. ■• 105. The skeleton, formed of polyhedral elements,
is coloured grey, and the soft tissue pink. The gelatinous layer of the upper surface is
seen to contain scattered cells (c. j.) and spherules. It forms a thin membrane (m.), im-
perforate in the portion seen in this section, over the spaces connected with the openings of
the canals. The beginning of the deeper-lying tissue with ciliated chambers (c. c/t.) is seen
at the lower part of the figure. To the right an embryo is seen lying near the mouth of a
canal.
Fig. 9. Section of an unsegmented egg showing the large nucleus (n.) and nucleolus.
X about 300.
Fig. 10. Section of a .segmented embryo. The stained centres of the skeletal elements
are seen in the tracts of decalcified skeleton about it. x about 300.
Plate XLVII.
Fig. 11. Part of a vertical section showing a pore (p.) in the surface membrane, x 105.
Fig. 12. Part of a vertical section through the interior, cutting two of the large
canals (c.) longitudinallj-. x 105. Small canals given off from the large ones are seen to
penetrate and divide in the tissues throughout which the ciliated chambers (c. ch.) are
scattered.
Fig. 13. Part of a thick transverse section, showing the network of skeletal canals in
the interior, and the unperforated cortical layer {cort.) of the skeleton.
Plate XLVIII.
Fig. 14. Part of a thin transverse section through the soft tissue of the interior, from
a decalcified specimen. The skeleton has been filled in in a diagrammatic manner, x 170.
Fig. 15. Part of a thin transverse section through the soft tissue of the interior, x 980.
The ciliated chambers are seen, with the flagellar processes projecting into them. Some larger
cells {I. c.) are scattered through the protoplasmic reticulum.
Fig. 16 ct and b. Two ciliated chambers, x 980. In h a small canal is seen to be in
close proximity^ to the chamber.
Fig. 17. Section showing communication between a ciliated chamber (c. ch.) and a small
canal (c'.).
Figs. 18 and 19. Groups of scattered cells in the jelly near the upper surface.
Fig. 20. Section through the soft tissue of a specimen whose canal system contains
masses of rod-shaped bodies {b.). x 250.
WiLLEY. Zoological Results.
Plate ZLVn
/kjC^r.
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y
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t-
.l..n.l?,t.-l d.:l
^/ -^tlu
i^^^
Wrjii , Nfwniiiri i^linmi'
t it.".ri':i'
WlLLEY ZOOLOGECAL ReSULTS.
c.ch^--
.^.
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! ■
i
>->^ J . .,1-,/*
7 ^%?
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. c ch.
rl.A.TZ
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't'^l^
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><
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16a X 980
18 ' 980
J.J. Lister del
V..- 'V'^
M^
m
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30. X 250
*^
■V'-
19.
LISTER ASTROSCLERA.
15. «980
■'Si'
let. X 980
17 "980
West, Hamnun liU-.
A CONTRIBUTION TOWARDS OUR KNOWLEDGE OF THE
PTERYLOGRAPHY OF THE MEGAPODII.
By W. p. PYCRAFT, A.L.S.
With Plate XLIX.
I. Introductory.
The Megapodes included iu Dr Arthur Willey's collection belong exclusively to
the genus Megapodius. It comprises a valuable series of embryos and one newly-
hatched nestling. To what species these belong it is not possible to say with certainty,
except iu the case of the newly-hatched chick, which is apparently M. eremita.
It has not been thought necessary to describe any-thing other than the pterylo-
logical characters of the specimens forming the subject of this Report, these being
hitherto almost unknown.
For the jjurposes of comparison, and for the sake of increasing the value of the
present paper, I have added a description — as complete as circumstances would allow —
of the pterylosis of the only adult preserved in spirit in our collection. This, being the
type of M. pritchardi, had to be examined so as not to inflict damage. Only the main
characters therefore of the adult pterylosis have been recorded ; from these, however, it
would appear that the differences between the adult and the embryo are but slight.
In so far as the description of the areas of the tracts is concerned it will be
found that the adult of M. pritchardi, and the embryo (p. 487) are the same.
Besides Dr Willey's embryos, the British Museum possesses one nearly ripe
Megacephalon maleu and a newlj'-hatched chick of Megapodius eremita, and these
have proved very helpful.
It is to be hoped that iu course of time the pterylosis of all the Megapodes
will be described. The present claims to be the most complete yet publi.shed.
II The Pterylosis of the Aoult {M. pritchardi).
Pterylae : —
Ft. capitis. This is a fairly evenly distributed tract, but the feathers of the
forehead, and sides of the head are very small and barely conceal the skin. The
inter-ramal area is not sharply defined. There are no eyelashes.
484 A CONTRIBUTION TOWARDS OUR KNOWLEDGE OF
Pt. colli. This divides about half-way down the neck into a pt. colli dorsalis
and ventralis. The former soon jjasses into the
Pt. spinalis. This tract is separable into two regions, an upper and a lower.
The upper, which represents a backward continuation of the pt. colli dorsalis, terminates
in a slightly bifurcated free end. The feathers of this region are much longer and
stronger than those of the remainder of the ti-act.
The lower moiety of the tract is of great width, and composed of long, rather
loose feathers. In the caudal region it becomes suddenly constricted and terminates
at the base of the uropygium.
Pt. caiidae. There are 12 rectrices. In the nestlings these are wanting.
Pt. colli ventralis. This does not appear, as a distinct tract, till about the
middle region of the neck, where it forms a pair of lateral bands ; these pass
downwards on either side into the
Pt. ventralis. This may be divided into a pair of antei'ior and external, and
jjosterior and internal tracts. The anterior are derived from a backward e.xtension,
and widening of the pt. colli ventralis. Each terminates near the posterior end of
the breast-bone. The posterior portion of the tract may be traced from the cloacal
aperture forwards as a narrow^ band of feathers running on either side of the middle
line, to within a short distance of the level of the clavicle.
Pt. ani. The bulk of the feathers of this small tract lie caudad of the cloacal
aperture.
Pt. femoralis. This is not a sharj^ly defined tract, but merges above into the
pt. spinalis and below into the pt. cruralis. The feathers along its inferior border
are much larger and stronger than those of the rest of the tract.
Pt. cruralis. This tract completely invests the leg, nearly as far downwards as
the tibio-tarsal joint.
Pt. humercdis. This tract is very sharjjh" defined, except anteriorly, where it
passes forwards to join the pt. ventralis.
Pt. alaris : — Metacarpo-digitals 10. Cubitals 1.5. Diastataxial '.
The first cubital is shorter than the second but not markedly so, as in the
Galli, e.g. G. bankiva The carpal remex, which should probably be regarded as the
real first cubital (4) is vestigial, plumaceous in character, and much smaller than its
covert.
Tectrices : —
T. viajores (dorsal). Those of the primaries are fairly large and strong ; of the
secondaries the chief point to be noticed is their relative length : unfortunately this
cannot be very certainly made out in the only adult available for examination,
' This term is synonymous with aquiutocubital.
THE PTERYLOGRAPHY OF THE MEGAPODII. 485
many of these and the neighbouring series having been cut. Enough remains to
show that the fifth covert is neither abruptly longer nor shorter than the sixth, as
sometimes happens in diastataxial wings (10).
The major coverts of the under surface of the wing are small.
T. mediae. On the dorsal aspect of the manus these feathers are very small ; they
run from the second remex outwards to the tip of the wing. It is interesting to
note that the sixth median cubital covert is markedly shorter than the fifth and
seventh — on either side — which two are of about equal length. The overlap appears
to be proximal.
This series appears to be absent on the ventral surface.
T. minores. One row of small, quite vestigial feathers on the manus, from 3-
rows of the cubital series.
The minor coverts of the ventral aspect of the wing are small and only imperfectly
clothe its surface.
T. marginales. Except along the extreme pre-axial border of the wing these
cannot be distinguished on the dorsal aspect from the minor coverts.
On the ventral surface they are readily separable with (1) those which clothe
the extreme pre-axial border, and point outwards, and (2) those which lie caudad of
these. The latter form a single row of elongated feathers pointing backwards, and
serving to clothe the patagial membrane. iioreover they entirely conceal the most
anterior of the minor coverts.
The parapteron is small, but does not otherwise offer any points of particular
interest.
The hypopteron ; like the parapteron this group of feathers does not call for any
special remark.
Ala spuria. This is of considerable size, the outermost and longest extends
backwards as far as the free border of the major coverts of the hand.
The carpus of the adult, from which this description is taken, is armed with a
thickened callous pad.
Semiplunue. A few occur — as usual — along the borders of the pt. spinalis.
Pluinuhe. These are very small, and confined to the pt. alans.
FilopltancB. One or two at the base of each of the larger contour feathers.
Bhamphotheca, simple, with a feebly-developed cere in which the nostrils are
placed. These last are impervious, and open, not protected by an operculum as iu
Gallinaj.
Podotheca bare, in the form of large plates.
Claws. Absent on the wing, on the foot very long and blunt.
Uropygimn, tufted.
Apteria : —
A. spinale. This is very small. In the embryo agreeing precisely with that of
G. banlciva, of about the same age.
w. IV. 65
486 A CONTRIBUTION TOWARDS OUR KNOWLEDGE OF
A. colli laterale. This divides the lower half of the pt. colli into dorsal and
ventral moieties (p. 484') ; it is a forward continuation of the
A. triuici laterale. The area of the space is much restricted by reason of the
great extension of the hinder portion of the pt. spinalis. It extends downwards and
backwards, so as to divide the pt. ventralis from the pt. cruralis, and is continued
forwards to the point where the pt. humeralis forms the pt. ventralis. Passing upwards,
above the wing and humeral tract, it becomes the Apt. colli laterale.
A. mesogastroei. This lies in the mid-ventral line and extends from the cloacal
aperture forwards to about the middle of the neck.
III. The NestliiNG Plumage.
Dr Gadow (5) coined the word Neossoptiles, to distinguish the plumage of
the nestling birds — when present — from the Teleoptiles or plumage of the adult.
Later, I showed that this nestling plumage was often made up of two different kinds
of down. The one preceding definitive down feathers, the other preceding definitive
contour feathers. For the former I proposed the term pre-plumulse, and for the latter
pre-pennse. Sometimes, as in the Duck, the nestling plumage consists of pre-penn£e
only, sometimes, as in the case of the young Hawk, of both kinds.
The nestling Megapode possesses no pre-plumulfe, and it is doubtful whether
pre-pennse, in the sense in which this word was originally used, occur. It seems
certain that the pre-penna3 representing the remiges and contour feathers generally
are shed during embryonic life (see p. 488). If this proves to be true, and there
seems to be room for but little doubt, we have a third form of nestling plumage —
a stage between the pre-penna proper, and the actual definitive feather of the adiilt.
In passing we might remark that this is unique only in so far as the time of its
appearance is concerned; for, as the wTiter has already pointed out (11), the young
Owl dons a special, woolly form of covering intermediate between the pre-pennse and
the definitive feathers, whilst, as already remarked, the pre-peima? of the Megapode are
lost in embryonic life, so that the young Megapode starts life in the same kind of
plumage as that assumed by the young Owl later on. The structure of this nestling
plumage is discussed on p. 488.
It is sufficient to remark here, that the nestling has no rectrices, and that
the development of the first cubital remex, as in the nestling chick, is arrested ;
its place being filled by a few downy filaments — probably representing a true neos-
soptile. The remex probably does not appear until after the first moult of the
" quilLs." The wing further agrees with that of the Common Fowl, and of
Opisthocomus — in that the development of the outer primaries is suppressed until
later in life. I have discussed the significance of this in recent papers (8, 10). It
is probable that in the case of the Megapode they do not, like the first cubital,
appear until after the first moult. In the nestling Megapode and Opisthocomus there
are 8 primaries, in the adult 10. In the nestling Chick there are 7 primaries, in the
THE PTERYLOGRAPHY OF THE MEGAPODII. 487
adult 10. There are no claws in the wing of the nestling Megapode, but they
appear for a while in the embryo.
The manus and fore-arm of the ? 6 — 7 day embryo ai-e sub-equal in length, in
the nestling the manus is considerably shorter than the fore-arm. This diftereuce or
change in relative length of the fore-arm and hand has been already commented
upon in the Common Fowl and in Opisthocomus, in the paper just referred to (10).
IV. The Pterylosis of the Embryo.
In so far as the general form and proportions of the tracts are concerned, the
embryo, as already remarked, tloes not differ from the adult. Furthermore, it is
difficult, given embryos of the same relative stage of development, to distinguish
between those of the Common Fowl and those of the Megapode. In the stage
figured, the two forms may be certainly distinguished by the fiict that, in the Fowl
the wing is eutaxial and there is a rudimentary " comb," and in the Megapode the
wing is diastataxial and there is no rudiment of a comb.
Concerning the diastataxial wing of this embryo (Fig. 4) I would remark only,
that it does not differ materially in the form and distribution of the shifting papillee,
from the normal diastataxial wing, as recentl}' described by me (12). There are no
claws on the wing of this embryo.
The wing of a much earlier embryo than this exhibits one or two features of
great interest, and, I think, importance.
In the first place, it appears to have already become diastataxial : the develop-
ment of the feather papillae is still very faint)}' traceable so that this point is
difiicult to make, nevertheless there is but little doubt that it is so. Furthermore,
the post-axial border of the distal end of the fore-arm has a swollen appearance, as
if the superficial growth had been faster than the deeper parts, so as to cause a
tendency which would ultimately end in producing a fold. It suggests that the rate
of gi-owth of the skin, and muscular tissue below, had outstripped that of the skeleton
withio. This has crowded the anterior cubital remiges together, and forced the
associated major coverts out of position, finally resulting in diastataxy. This swollen
stage is a noticeable feature in many diastataxial wings. I do not, however, in the
above facts see any reason to alter my recent explanation (12) as to the manner in
which diastataxy is brought about. I should regard them rather as secondary phenomena
probably not connected with the origin of diastataxy at all.
The length of the hand in this stage is less than that of the fore-arm.
This collection contains several embryos much younger than the last, and two of
stages much older than that here figured (Fig. 1). With the very early stages we
cannot deal in the present paper ; the later stages are useful — in the present connection —
only in so far as they afi'ord material for studying the nature and development of
the nestling plumage, and will be dealt with under this head.
65—2
488 A CX3NTRIBUTI0N TOWARDS OUR KNOWLEDGE OF
V. Structure of the Feathers, axd the nature of the Nestling Plumage.
Contour feathers : the structure of the adult contour feathers calls for no special
remark, all the feathers of the trunk possessing a normal, well-knit, coherent vexillura.
This fact, however, afifords a valuable and important standard of comparison between
the contour feathers of the adult and those of the nestling, and brings out one very
interesting point of difference between these two stages.
The plumage of the newly-hatched Megapode will probably prove to be unique.
It differs from the definitive contour feathers of the adult on the one hand, and the
typical pre-pennffi, which precede these, on the other. It agrees very closely ^\-ith
that recently described by me (11) in the nestling Owl; in that, like this, the trunk
feathers are of a loose, semiplumous character, the vesillum of the feathers being
disconnected.
By a typical pre-penna, I should e.-cplain, that I take as my standard of comparison
the pre-penna of the strictly Gallinaceous birds, e.g. Turkey (PI. XLIX, Fig. .5). In
this we have, microscopically examined, a distinct main-shaft, and an after-shaft, each
provided with rami and radii; the latter bearing feeble fila (Fig. 11 F.). The
after-shaft breaks up at once into rami, whilst in the main-shaft the rhachis is
traceable upwards for a considerable distance.
A feather, taken from the same region of the body of a ripe embryo or of a
newly-hatched Megapode chick, shows a distinct advance upon this. As will be seen
in the figure (Fig. 6) the main-shaft is long and tapering, and bears numerous
bilaterally arranged rami ; the after-shaft likewise is well defined, and bears rami
disposed as in the main-shaft Furthermore, the rami and radii differ slightly in
structure.
In the Turkey the rami were produced into fine terminal filaments beyond the
last pair of radii, and the radii were seen to be provided only with feebly developed
fila; this was the case no matter what part of the feather was examined.
In the Megapode the rami and radii, as in the more perfect contour feathers,
differ somewhat in structure according to whether those examined are at the base
or the tip of the feather. The rami from the base are long, and slender, and
the radii take the form of long delicate rods bearing more or less well-marked nodal
swellings or flanges at intervals (Fig. 10), which may be produced forwards into fila. The
rami, from the distal end — of the main-shaft only — resemble rather those of the pre-
pennae of the young Tinamou which I described some time since (11), inasmuch as each
ramus is laterally compressed at the base into a blade-like lamina, which, tapering
forwards, gradually becomes segmented and rod-shaped; the fila are here constant, paired,
and proceed from the distal end of each segment. These fila are the homologues
of the " booklets " of the more perfectly developed contour feathers : indeed, booklets
occur, though feeble, in the feather from which this description is taken. The nestling
down feather of the Megapode, however, differs from that of the Tinamou chiefly in
that the rami are somewhat weaker.
In spite of this resemblance between the nestling down plumage of the Tinamou
and the Megapode, there can be no doubt that the feathers composing each differ in
THE PTERYLOGEAPHY OF THE MEGAPODII. 489
one important point. The former are true pre-pennse, the latter are somethinjr rliffor-
ing from, and intermediate between, pre-pennte on the one hand and definitive
contour feathers on the other. They represent apparently the adolescent plumage
which we noticed was assumed in the case of the Owls (12) during their first year;
but with this difference : in the Owl, these semiplumous feathers were preceded by
true neossoptiles, in the shape of pre-pennse ; in the young Megapode they are not :
for these feathers are probably only to be found ujion the embryo, as will be seen below.
The above facts raise a most interesting question concerning the life history of
the Megapodes. For it may now be asked : If the plumage of the young Megapode
is not made up of Neossoptiles — or pre-pennse and pre-plumulte in the strict sense of
the word — but of a something intermediate between pre-pennsje and the definitive
contour feathers of the adult, is the development of these neossoptiles entirely sup-
pressed or rather entirely eliminated ontogenetically ?
The material represented in this collection will I think afford an answer to this
question, but more specimens, and in a slightly better state of preservation, will be
necessaiy before it becomes absolutely convincing. For my own part, I have no
doubt at all, and would reply that the development of pre-pennse has been almost
totally eliminated, though transitory rudiments still make their appearance.
These can well be studied in the ripe embryo, in which they appear as long
straw-coloured filaments depending from the tips of tiie remiges.
Under the microscope, these filamentous processes prove to be the sheath of the
outer Malpighian layer, which invests the developing feather. Through this the
rudiments of rami can be plainly seen. These are apparently quickly disintegrated, but
the capsular sheath remains unbroken, until thrust out, upon the tip of the definitive
feather replacing it, and from which it is ultimately cast off. Normally, this sheath
would have been broken up and dispersed by the developing rami of the pre-penna,
which, later, would have been carried out on the tip of the definitive feather
destined to replace it. In the present case, as we have just seen, the rami never
attain strength — or bulk — sufficient to burst this investing sheath, which remains, for
a season, in the place of the pre-penna dissolved within it, attached to the tip of
the definitive feather, which has superseded it.
In younger embryos, in that stage in which the feathers look not unlike much
elongated Porcupine quills, it will be noticed that each of these "quills" becomes
slightly constricted near the tip, swelling out again distad of the constriction, and
then tapering rapidly to a point (Fig. 7). I take it that this constriction occurs at
what was once the base of a pre-penna, and that the swollen, distal end of this
" quill " represents all that now remains of it. A comparison of Figs. 7 — 8 will make
this clearer. Fig. 7 represents the " quill " stage just referred to ; in this the
rami of the definitive feather seem clearly distinguishable from the fine rami that I
take to be remnants of pre-pennse. Fig. 8 is a later stage, drawn from one of
the "straw-coloured" filaments from the tip of a remex. Fig. 9 shows the pre-penna
of a Turkey attached to the distal end of the rami of a definitive feather.
One of the most striking features of the ripe embryo, whereby it is seen to
differ at once from all known birds, is the great length of the remiges, and their
490 A CONTRIBUTION TOWARDS OUR KNOWLEDGE OF
coverts, which lie on either side of the body like a series of long closely packed
camel-hair pencils — the tips of the feathers having already burst the investing sheath
in which they were enveloped — extending backwards to the extreme end of the body.
It is in the stage earlier than this, before the tip of the sheath has burst, that the
rudimentary pre-pennse are found.
Summary.
As stated in the earlier part of this paper, this is the first attempt at anj'thing
like an extended or detailed account of the Pterylography of the Megapodes, and even
this leaves much still to be done.
The only other record of the pterylological characters of this group is that of
Nitzsch (6). He was perforce obliged to content himself with an examination of a
stuffed specimen. In spite of the difficulty of such an unsatisfactory method of
investigation, he managed to make out enough to show him " it possesses the general
characters of the Gallinacea;," and that the oil-gland was tufted.
Since then the additions to our knowledge of this subject have been few, and
chiefly concern the fact that the young chick is enabled to fly almost immediately
after leaving the egg.
The most important of these is that of Dr Gadow (2), who ^vrites : " Die Jungen,
wenigstens einiger Arten, schliipfen vollstiindig befiedert und flugfahig aus ; das Xestkleid
ist also embryonal unterdrllckt worden."
Dr Sclater (13) was apparently the first to point out, on the authority of Mr
Beddard, that the wing of at least one species of Megapode (M. rubrifrons = ereniita)
was " aquintocubital " = diastataxial.
The more or less precise details of the pterylological characters of the adult and
embryo given in the present contribution will it is hoped be shortly supplemented
by similar descriptions of other genera. The facts embodied in this part of the paper
are really all new. Again, no one appears to have previously noticed the facts concern-
ing the ai-rested development of the outermost primaries, and of the 1st (2nd) cubital
remex in the nestling, or the differences in the rate of growth of the manus and
fore-arm. All these points, it is interesting to notice, the young Megapode shares
with the young Gallus and Opisthocomus.
The history of the pre-penna3 still needs further investigation. That the nestling
was not clothed in a covering of true nestling down has been more or less accepted
as a fact for a long while, but the difference between this nestling plumage and
definitive feathers seems not to have been previously remarked.
One or two problems concerning the development of feathers generally have been
raised during the present investigation. These however do not concern the questions
of pterylosis at all, and will be dealt with in due couree.
THE PTERYLOGRAPHY OF THE MEGAPODII. 491
REFERENCES.
1. Gadow, H. Bronn's Klass. u. Ordnung des Tliier-ieichs. Bd. 6, Abt. iv. Anatomisch.
Theil, 1891, p. 699.
2. Gadow, H. Ibid. Systemat. Theil, 1893, p. 171.
3. Grant, W. R. O. Catal. Birds Brit. Mus., Vol. xxii., 1893, p. 445.
4. i\TlTCHELL, P. C. " On so-called ' Quintocubitalism ' in the Wing of Birds ; with speei.al
reference to the Columbte, and Notes on Anatomy." .Journ. Linn. See. (Zool.), Vol.
XXVII., pp. 210—36, 1899.
5. Newton, A. Dictionary of Birds, 1896, p. 243. [Article " Feathers" by H. Gadow.]
6. NiTZscH, C. L. " Pterylography."' English Trans. Roy. Soc, p. 811, 1867.
7. Pycraft, W. p. "The Interlocking of the Barbs of Feathers." Natural Science, Vol. 3,
1893, pp. 197—203.
8. Pycraft, W. P. "The Wing of Archreopteryx." Natural Science, Vol. v., 1894,
pp. 3.50—60.
9. Pycraft, W'. P. "A Contribution to the Pterylography of the Tinaniformes." Ibis,
1895, pp. 1—21.
10. Pycraft, W. P. "On the Pterylography of the Hoatzin." Ibis, 1895, pp. 345—73.
11. Pycraft, W. P. "A Contribution towards our Knowledge of the Morphology of the
Owls." Trans. Linn. Soc, Vol. vii. (2nd Ser. Zool.), 1898, pp. 223—275.
12. Pycraft, W. P. "Some facts concerning the so-called ' Aquintocubitalism ' in the Bird's
W^ing." Journ. Linn. Soc. (Zoology), Vol. xxvii., pp. 236—56. 1899.
13. Sclater, p. L. "Remarks on the Fifth Cubital Reme.x of the Wing in the Carinatae."
Ibis, 1890, pp. 77—83.
14. Wallace, A. R. "Malay Archipelago," p. 305. 1890.
EXPLANATION OF PLATE XLIX.
Fig. 1. Lateral view of the left side of an embryo Megapodius eremita, showing the
form of the pterylfe. The dotted areas represent pterylte ; the relative sizes of the feathers
are indicated by large and small dots.
Apt. coll. hit. Apterium colli laterale.
Ajjt. m. Apterium mesogastroei.
Pt. cap. Pteryla capitis.
Pt. caud. Pteryla caudalis.
Pt. coll. dors. Pteryla colli dorsalis.
Pt. coll. vent. Pteryla colli ventralis.
Pt. cr. Pteryla cruralis.
Pt. f. Pteryla femoralis.
]'(. hum. Pteryla humeralis.
Pt. .y). Pteryla spinalis.
Fig. 2. Dorsal view of the same. The dotted parts as before.
Apt. sp. Apterium spinale.
Apt. t. lat. Apterium trunci laterale.
Fig. 3. Ventral view of the same.
492 A COXTRIBUTION TOWARDS THE PTERYLOGRAPHY OF THE MEGAPODII.
Fig. 4. Dorsal aspect of right wing of embryo (Fig. 1) showing the first appearance
of the feather-papillaj representing the remiges and major coverts of the fore-arm. The
arrangement is seen to be diastataxial.
c. c. Carpal covert.
c. r. Carpal remex.
i. c. Intercalary coverts.
med. c. 3Iedian coverts.
mj. c. Major coverts.
min. c. Minor coverts.
Ith c. r. 7th cubital remex.
Fig. 5. A pre-penna of a nestling Turkey. To show the after-shaft, and widely sepa-
rated radii. Note the filamentous ends of the rami.
A. After-shaft.
D. f. Definitive feather.
Rd. Radius.
Jivi. Ramus.
Fig. 6. A feather from a nestling Megapode. This should be contrasted with Fig. .5.
Fig. 7. Portion of a definitive feather supporting a rudimentary pre-penna from a late
embryonic stage of Megapodius eremiia.
D. f. Definitive feather.
P. Pith cells.
P.p. Pre-penna.
R. Rami.
Fig. S. Portion of a definitive feather with rudimentary pre-penna from a ripe embryo
of same species as Fig. 7.
Fig. 9. Portion of a definitive feather of a nestling Turkey, supporting a pre-penna,
the distal portion of which has been removed. The rami of the pre-penna have burst the
investing sheath and are now free.
Fig. 10. Portion of a radius from the base of a feather from a nestling Megapode.
Fig. 11. Portion of a radius from the pre-penna of a nestling Turkey, i^. = Fila.
WlLLEY ZoOliOGICAL, RBStTLTS.
Plate XLIX.
pt.t.z<u ^1* 11/5, a'
■W.PPycraft.dd. admt
West,Ncwinan lith.
PYCRAFT. MEGAPODIUS.
THE STOLONIFERA AND ALCYONACEA COLLECTED BY
DR WILLEY IN NEW BRITAIN, ETC.
By SYDNEY J. HICKSON, F.RS.,
Beyer Professor of Zoology in the Owens College,
AND ISA L. HILES, B.Sc,
Owens College.
With Plates L. and LI.
STOLONIFERA.
1. Tubipora inusica, L.
A single specimen well preserved in spirit of this widely distributed species was
sent to us. It was obtained at Welle Island, D'Entrecasteaux group. Attention has
already been called by one of us (7) to the unsatisfactory nature of the evidence
for dividing the genus into a number of species, and the comparison of both hard
and soft parts of this specimen with specimens obtained by Dr Gulliver in Zanzibar
and by one of us in Celebes confirms the opinion previously expressed that there
is only one species of the genus. The principal character which has been used for
distinguishing species is the diameter of the polyp walls, but this character in every
specimen depends entirely upon the situation on the reef in which it happened to
grow. It is what would be called in modern phraseology " an acquired character," and
there is no evidence that it is one which is inherited as all true specific characters
may be. The tubes happen to be in this specimen 2'5 mm. in diameter, but there
can be little doubt that if Dr Willey had collected specimens from different localities
great variations would have been found in this character.
If we accepted the current views on the species of Tubipora, Dr Willey's specimen
would be called T. rubeola Q and G (20). In describing their new species these
authors say that compared with T. viusica " les tubes sont plus gros, plus long, legers,
moins serrds et offrant cylindres de deux a trois pouces sans nojuds." It is true
that the distance between the platforms does not amount to more than 10 mm., but
the specimen is a young one and in other respects resembles the description of T.
rubeola Q and G.
w. IV. ()6
494 THE STOLOXIFERA AND ALCYONACEA
The anatomy of Tuhipora has already been fully described (12 and 8) but it
seemed desirable, since Dr Ashworth (1) has shown that there is a marked divergence
from the general anatomical features of the Alcyonarian stomodaeum and mesenterial
filaments in the genus Xenia, to reinvestigate the structure of these organs in
Tuhipora.
The stomodaeum possesses an ectodermal epithelium of clear columnar cells as
in Alcyonium and other forms, and there is a well-marked siphonogh-ph armed with
long cilia. No gland cells are found in this epithelium. There are six short ventral
mesenterial filaments bearing gland cells. These filaments begin on the free edge of
the mesenteries in contact with the inner opening of the stomodaeum. In the con-
tracted spirit specimens they may be seen to extend about 5 mm. down the free
edges of the mesenteries. It is impossible to estimate what their exact length may
be in the living expanded polyps. The two dorsal mesenterial filaments are very
much convoluted and extend from the stomodaeum as far down as the first tabula.
There are no gland cells in their epithelium.
Judging from these anatomical facts then, the digestive processes of Tuhipora
must be carried on as in Alcyonium and Kophohelemnon, the digestive fluid being
secreted by the six ventral mesenteries and not in any degree by the epithelium of
the stomodaeum nor by the epithelium of the dorsal mesenterial filaments.
The specimen was a male. The young sperm sacs in their follicles were found
on the six ventral mesenteries only, as described and figured by von Koch\
2. Glavularia viridis Q and G.
A small piece of a colony of this species was sent to us in a partially retracted
condition. It was obtained at Welle Island (Sanaroa), D'Entrecasteaux group, in 1895.
The specimen differs from those obtained in Celebes (9) in being decidedly
stouter in build. The polyps are not very long (about 35 mm.), springing from a
creeping stolon attached to a piece of dead coral. The terminal extremities of the
polyps have an inflated appearance due, undoubtedly, to the manner in which they
were killed. They are from 40 — 50 mm. in length and 8 mm. in diameter at the
free end. Only one transverse tubular connection between the polj'ps above the level
of the stolon was seen.
The spicules vary considerably in length but the average is about 2'5 mm.
The tubercles are rather less prominent than those of the spicule figured by one of
us (9), but similar to those of other spicules in the collection from Celebes.
In the larger spicules of Dr Willey's specimen there may be frequently seen a
curious bifurcation of one end. The same has been noticed in the Celebes specimens.
The lower, and therefore older, parts of the body wall are strengthened by the forma-
tion of horny fibres in the mesogloea, making the texture hard and brittle.
' On referring to my preparations made in 1884 I find that 'I was in error in the statement I made
then that the gonads occur on the dorsal mesenteries in the specimen from Zanzibar.
It is extremely probable that Dana (2) was mistaken in stating that the specimen he examined was
hermaphrodite, six of the mesenteries bearing sperms and two ova. Hermaphroditism is of very rare occurrence
in Alcyonarians and the presence of gonads on the dorsal mesenteries has not yet been satisfactorily proved
in any genus. S. J. H.
COLLECTED BY DR WILLEY IN NEW BRITAIN, ETC. 495
The dorsal mesenterial filaments extend from the free edge of the stomodaeum
to the stolon. The}- are considerably convoluted and drawn out laterally into long-
tag-like processes (Fig. 14). The six ventral mesenterial filaments are short, beginning
some distance from the free edge of the stomodaeum as shown in Fig. 13. The
gonads (the specimen was a female) occur below the ventral mesenterial filaments
and are attached to all six of the ventral mesenteries. Numerous nematocysts
(001 mm.) were observed in the peristome. They are spindle-shaped, with very fine
points.
There can be no doubt that Dr Willey's specimen belongs to the same species
as that originall}' described by Quoy and Gaimard from Vanikoro, as the specimen
obtained from the Aru islands by Wallace (now in the British Museum) and as those
which occur on the reefs of Celebes. It is very similar in some respects to the
specimen described as Clavidaria injlata by Schenk (24), and it is quite probable that
Schenk's specimen was but a young form of this species.
Now that we know something more of its anatomy it may seem to be an open
question whether it would not be advisable to constitute a new genus for this species,
the principal characters of which would be the tubular connections between the body
walls of adjacent polyps in large specimens, the horny fibres of the mesogloea, the
large spicules, the tag-like processes of the dorsal mesenterial filaments and the remote
position of the ventral mesenterial filaments. We do not propose however to give it
a new generic name in this paper, although we wish to emphasise the fact that the
species stands very much apart in the genus. It is a matter of some regret that
owing to the manner in which the polyps contracted when killed they are not suffi-
ciently well preserved to give good results of cellular histology. We are not able
therefore to say for certain whether gland cells occur in the dorsal mesenterial filaments
and in the stomodaeum or do not. All that can be said is that the histology is
apparently very different from that of Tubipora and that the matter is well worthy
of further investigation. It may be mentioned for the guidance of collectors that
when polyps exhibit an inflated appearance after death they are usually distended
with sea-water and practically sealed to the preserving fluid in which they are
immersed. Such polyps should be slit open and placed in fresh spirit or the endoderm
will macerate and the specimens be useless for histological investigatiou.
ALCYONACEA
The specimens collected by Dr Willey, which belong to this sub-order, have proved
to be of considerable interest.
There are two species of the genus Telesto, one of which is new, two species of
Nephthya, and five species of Spotigodes, one of which is new.
The sub-family Siphonogorginae is represented by one species of the little known
genus Chironephthya.
The Alcyonidae are represented \>y four species of Alcyonium, of which one is
new, three species of Sarcophytum and four of Lohophytum.
The representatives of the Xeniidae are five in number, one being new to science.
They are described in a separate paper by Dr Ashworth.
66—2
496 THE STOLONIFERA AND ALCYONACEA
3. Telesto rupicola, Miiller (PL L., Figs. 1, 2).
There is one specimen of this species, the upper portion of which is thickly
covered by a commensal sponge through which only the polyp's tentacles protrude.
The branches of the basal stolon are from 1-5 — 20 mm. in length by 1 mm. in
diameter.
There is one main axial polyp, iiO mm. long by 2 mm. in diameter. It bears
three secondary axial polyps, all on the same side ; the first 18 mm. long is borne
52 mm. from the base of the axial poh"p ; 64 mm. above this is the second, 8 mm.
long; the third is 8 mm. above the second, and is 16 mm. long.
The axial Tpolyp also bears lateral polyps at intervals of from 2 — 4 mm.; they
are irregularly placed round the axis. The stolon bears several independent polyps,
and the secondary axial polyps bear tertiary polyps (Fig. 1).
The expansible portion of the polyps is 5 mm. long (the tentacles being 3 mm.
long)'. There are no spicules in the tentacles but there are eight small groups of
a few narrow spindles extending from the bases of the tentacles a short distance
down the polyp. The body wall of the non-retractile part of the polyp is filled
with spicules. All the polyps are expanded in the specimen. The primary and
secondary axial polyps are of the same type. The cal}'x walls are marked by eight
longitudinal grooves.
The spicules are long, narrow, spiny spindles, often branching at one end, thus
becoming Y shaped (Fig. 2).
The spiny spicules are "607 mm. x "11 mm. — '22 mm. x '092 mm. (including the
spines) ; the branched forms are : — the main stem "368 mm. by '092 mm., one branch
'092 mm., the other "184 mm.; from the tip of one branch to that of the other,
•276 mm.
The spicules are colourless. The colony is brownish-white in colour in spirit.
Habitat. Blanche Bay, New Britain. Depth, .50 fathoms. Previously recorded from
the coast of Brazil, and Bahia (19).
4. Telesto arthuri, sp. nov. (PI. L. Figs. 3, 4. PI. LI. Fig. 12).
A single specimen, incomplete at the base, differs in such a marked degree from
any species of the genus that bas been described that we have found it necessary
to constitute for it a new species, which we propose to name after its discoverer,
Dr Arthur Willey.
There are three branches in the specimen (Fig. 3), the longest being 55 mm. in
length and 4 mm. in diameter. Numerous polyps contribute to the substance of each
branch ; the difference in this respect between our new species and Telesto rupicola
being very striking, as ma}- be seen by comparing Figs. 1 and 3. The portions of
the polyps which project from the branch are arranged in an irregular spiral, and
each one inclines to the branch at an angle of 45°. • The projecting portion of each
' The necessity for the introduction into the literature of the Alcyonaria of some term to express that
part of the polyp which can be expanded and retracted is obvious. Mr G. C. Bourne has proposed in a paper
read before the Linuaean Society to call it the " Anthocodia," but as Mr Bourne's paper is not yet published
we do not intend to adopt it here.
COLLECTED BY DR WILLEY IX NEW BRITAIN, ETC. 497
polyp consists of two parts ; a part which can be completely retracted and a part
which owing to the constitution of the wall cannot be retracted. In the specimen
the first named part of each polyp was retracted, and we cannot give any estimate
of its length in consequence. The non-retractile portion of each polyp however pro-
jects about 2 mm. from the branch and is about 1 mm. in diameter. The substance
of the branches is fairly soft, the coenenehyni being leathery in consistency, strength-
eued at its periphery by long, minutely warted spindle-shaped spicules (Fig. 4). These
spicules vary from 3'3 mm. x '27 mm. to l.S mnj. x 'IS mm.
Habitat. Blanche Bay, 30 fathoms.
This species differs from all the other species of the genus which have been
adequately described, in the size and character of the spindle-shaped spicules and in
the feature that the polyps are numerous and crowded The absence of longitudinal
grooves distinguishes it from Telesto riisei (Duch. and Mich.) and T. arborea (W. and S.).
Anatomical structure (PI. LI. Fig. 12). On making a transverse section through
one of the branches, it may be seen to be perforated by a number (8 — 10) of polyp
cavities arranged in a d-efinite manner as regards the axis — the dorsal side of each
polyp, indicated by the dorsal mesenterial filaments, being turned towards the axis
as in other Alcyonarian colonies. The raesogloea is moderately thick and is penetrated
by numerous endodermal canals (c.c). These canals are in all cases of small diameter,
no large endodermal canals such as we find in Xema (1) and in some of the Gor-
gonacea being found in this genus. The canals communicate with the polyp cavities,
and from the fact that the polyp cavities communicate with these canals at their
proximal ends, as they do in Alci/oniuui, it is probable that they give rise to the
young buds. The polyp cavities do not communicate with one another directly. The
substance of the mesogloea also contains a number of small bipolar cells, but the
preservation of the mesogloea is not iii a sufficiently satisfactory state to allow us
to follow the histology into further details.
It can be positively stated that there is no axis of any kind, not any axial
canal comparable to the axial canal of Coelogoiyia. Notwithstanding the resemblance
to a Gorgonacean, therefore, that this species of Telesto may be considered to have,
it is unquestionably Alcyonaceau. The specimen is a female but the gonads are
probably very immature.
Family. Xephthyidae.
This family is represented in Dr Willey's collection by several specimens which
belong to the genei-a Spongodes and Nephthya.
The genus Spongodes appears to be fairly well-defined and is widely distributed
in the shallow waters of the tropics of the Old World. A great many species have
been described based upon such characters as size, colour, form and the character
of the spicules. Many of these so-called species appear to be very closely related,
and it is po.ssible that on a revision of the genus based upon anatomical details
many of the specific names will have to be struck out of the list. The genus
Nepldhya presents us with many difficulties, and not the least of these is that some
498 THE STOLONIFERA AND ALCYONACEA
authors still separate the species without supporting bundles of spicules in the calices
from those with such bundles, into a separate genus called Ammothea.
This is one of several instances in the Alcyonaria of genera based solely on the
character of spicules, and considering the well-known range of variation in both the
number and shape of spicules in individual species of Alcyonarians, it is most desirable
that such generic names should be discountenanced.
But the name Ammothea ought to disappear altogether from the Class, on the
gi-ound that it was introduced by Leach in 1814 for a Pycnogonid (and is still used
in that sense) before it was used for an Alcyonarian. It is really remarkable that
the authors of the Challenger volume on Alcyonaria, knowing this fact, should still
retain the term Ammothea for an Alcyonid. It is difficult to see how such a course
can be defended.
Sub-family I. Spongodinae.
5. Spongodes cervicornis, Wright and Studer.
This specimen agi-ees in general form and method of branching with the type
specimen (28). The aiTangement and size of the spicules is also the same. They
are 4"3 x '18 mm., .5 5 x SI mm. — '73 x 03 mm., 1 x '018 mm.
The polyp heads are '8 mm. long. The colour of the main stem, branches and
polyp heads is whitish, the twigs are purple.
Habitat. Sandal Bay, Lifu.
Previously recorded from Tahiti.
6. Spongodes hemprichi, Klunzinger.
There are two specimens of this form, the larger of which is 72 mm. in height ;
the stalk is 3-5 mm. high and 20 mm. wide. The spicules on the branches are large,
opaque, white spindles covered with compound-tuberculate warts. These spindles are
transversely placed and are 48 mm. by •.5-5 mm. in diameter.
The polyp heads are in clusters; each has a spicule projecting above the head.
These spicules resemble those of the branches. Some of the spicules of the stalk
show a tendency to branching. They are 1'9 mm. long x '33 mm. — 'SS x "07 mm.
The colony is brownish-drab in colour ; the polj^D heads are brown. Some of the
spicules of the heads show a tendency to brown coloration, but the majority are white.
Habitat. Sandal Bay, Lifu.
Previously recorded fi-om the Red Sea (11).
7. Spongodes rhodosticta, Wright and Studer.
There is one specimen of this species.
The total height of the colony is 90 mm. The length of the stalk is 25 mm.
and its width is 3 mm.
The branches are on an average 10 mm. long. The poh^s are 2 — 8 mm. in
length, and the heads 1 mm. in diameter.
Habitat. Talili Bay, New Britain.
Previously recorded from the Arafura Sea (28).
COLLECTED BY DR WILLEY IN NEW BRITAIN, ETC. 499
8. Spongodes semperi, Studer (PI. L. Figs. 5, 6, 7).
There are several fragments and one small colony of this species.
The stalk is thick and cylindrical, widening out a little at the top.
Fi-om the upper end it gives off a ring of sis conical branches, two of which each
bear a small branch. The branches stand out almost horizontally, so that they form
a shallow cup at the top of the stalk.
Each is covered with closely set, .spirally arranged, fairly large polyp heads which
are surmounted by one or two long spicules. The branches ac(iuire the form of spikes.
The trunk is 11mm. high and 11 mm. wide. The diameter of the crown of
branches is 26 mm. The branches are 10-14 mm. long, by 3"5 — 4 mm. broad.
The whole colony is rigid ; the walls of the trunk are filled with long, thin,
slightly curved spindles closely covered with small warts and variously placed. The
same spicules are found in the branches and polyps, mingled in the latter with smaller
ones of the same type.
In some cases one, and in others two, of the long spicules extend throughout
the length of the pol\-p and project about 1 mm. beyond the head. The spicules are
from 4 mm. long by 02 mm. broad, to '06 mm. long by '00.5 mm. broad.
The colour in spirit is white.
This form agrees fairly closely with Studer's (25) description of Spongodes semperi.
The points of difference are the absence of the polyp-covered circular fold described
by Studer and of branches in the middle of the head.
Habitat. Sandal Bay, Lifu. Depth 30 fathoms.
Previously recorded from the Pliilippine Islands (2.5).
9. Spongodes rakaiyae, n. sp. (PI. LI. Fig. 11).
According to Kiikenthal's classification (17) of the genus Spongodes this belongs
to the third group Divarkutae, sub-group Cijlindratae, in which sub-group he includes
seven species. Of the seven species the form to which this specimen seems most
closely related is Spongodes klunzingeri Studer {Sp. ramulosa Klz.).
The colony is 200mm. high, and 130mm. wide; the main stem is 23mm. thick;
the primary branches are from 30 — 95 mm. long; the twigs are 12-20 mm. long.
The stem is free from branches for 35 mm. above the base, at which point it
bears a thick branch 70mm. long; 20mm. above this branch arise two large branches,
one on each side, one 70 mm. long, the other 95 mm. long. From that point the
stem bears several smaller branches from 30— 70 mm. long. For 75 mm. from the tip
the stem bears twigs covered with polyps. Below that point the stem is quite free
from polyps, which are borne on twigs springing from the branches only.
Each twig bears from 15 to 30 polyps. The twigs arise close together on all
sides of the branches and gradually get smaller towards the tips of the branches, so
that the latter have a conical shape. The jKilyp heads are not closely pressed against
the twigs but each has a short peduncle, and as each polyp stands out stiffly owing
to the bundle of long spicules supporting it, one of which projects for a short
distance ("5 mm.) beyond the polyp head, the twigs have very much the appearance of
small thorny branches.
500 THE STOLONIFERA AND ALCYOJSTACEA
The polyp peduncles are about 1-2 mm. long, the heads are '7 mm. high and •6 mm.
in diameter.
The spicules of the polyps are slender, spindle-shaped, and covered with fine small
spines. They are 'SI — 'llmm. long by 'OS — "01mm. broad. Small rod-like spicules
are arranged transversely in two iiTegular rows up the back of the tentacles for their
whole length.
The spicules which form the bundle supporting the polyp head are long, spindle-
shaped, and covered with small spines. They are from 1 to 2 mm. in length.
The upper portion of the stalk is loosely covered with large, warted, spindle-
shaped spicules which lie in all directions on the surface. They are 2 to 3'8 mm.
long and IS to "21 mm. broad.
The lower portion of the stalk contains a number of in-egular, knobbed spicules
covered with warts. They are '42 — ■2 mm. long and '18 — ■07 mm. wide. The colony is
pale fawn colour in spirit. The spicules are colourless.
Habitat. Straits of Rakaiya, Blanche Bay, New Britain. Depth, 3 — 4 fathoms.
The species to which this form seems most closely allied is Spongodes klunzin-
geri Studer (25) {Sp. ramulosa Klz.). It differs from it however in one or two points.
In Sp. hlunzingeri the spicules of the polyps cause them to be of a red colour.
In this form the spicules are colourless. The colony is much more branched and the
twigs also are more divided than in Sp. klunzingen.
The sjjicules of the polyp head are not so regularly arranged en chevron as sho\Mi
by Klunzinger in his figure (Taf. III. Fig. 2) and do not project beyond the re-
tracted tentacles. The spicules of the upper portion of the stem are smaller than
in Sp. hlunzingeri, where they are 3 — 5 mm. long. Also the spicules are all more
slender and regular than Klunzinger figures them. Consequently it seems best to call
this form a new species.
10. Nephthya chahrolii, Klunzinger.
There is one fairly complete colony w-ith two fragments.
The colony is 45 mm. high and 70 mm. b}' 40 mm. wide. The stalk is 20 mm.
high. It divides into four main branches, which bear numerous lateral branches and
these may again divide into branchlets. The ends of the branchlets are crowded with
round poh*}3 heads and look like short blunt spikes. The large branches are 20 — 30 mm.
long and 15 — 30 mm. wide. The branchlets are 2 — 10 mm. long, 4 — 8 mm. wide. The
heads are 1 mm. high.
The spicules agree closely in form and size with Klunzinger 's species (11).
The colour is pale fawn. The colony is female, but the ova are immature.
Habitat. China Straits, British New Guinea.
Previously recorded from the Red Sea.
11. Nephthya virescens, Sav.
This form belongs to the section of the genus called Ammothea by Savigny and
other authors, as there is no definite " Stiitzbundel" and the polyps are arranged in
" catkins." There are three large specimens of this form, two complete, one -with the
basal part missing.
COLLECTED BY DR WILLEY IN NEW BRITAIN, ETC. 501
The larger of the two complete specimens is 22o mm. high and 90 mm. wide.
The main stalk is 20 mm. in diameter at the base. The extreme basal part is
fairly rigid but the rest of the colony is soft and flabby. There are three branches
springing from the base which form small stalks of the same type as the main stalk.
The main stalk soon gives off various small branches ranging in length from 10 — 50 mm.
These bear small branchlets on which the polyps are so closely crowded as to
form spikes. At a distance of 120 mm. from the base is one large branch 70 mm.
in length which gives off several secondary branches about 1.5 — Sri mm. long. The tip
of the stalk is crowded with small spike-like branches.
One of the colonies has the basal part missing and is more branched than the
other two, the main stalk giving off three large branches which reach to the top of
the colony and bear numerous small branchlets covered with polyps.
There are no polyps borne directly on the main stalk or on the jjrimary branches ;
they are all borne in closely crowded spikes on secondary small branches.
The spikes are from 7 — 10 mm. long. The polyps are non-retractile but bend in
towards the branch on which they are borne. They are 1 mm. long ; the heads are
"5 mm. in diameter.
The polyps contain a few irregularly placed spicules, spindle-shaped and bearing
a few small spines which have a tendency to become more numerous and larger at
the ends of the spicules. They are '6 — 16 mm. long and 03 — '01 mm. broad. The
tentacles contain two irregular rows of transversely placed small rod-like spicules.
The stalk contains a number of spindle-shaped spicules covered with fairly large
conical warts ; some of these spicules are slightly curved and the warts are decidedly
larger on the convex side.
These spicules are 1'02 — '18 mm. long and "18 — '09 mm. broad.
The stalk also contains some small, four-rayed forms and some very rough, knobbed
forms with a small constriction in the middle, so that they look like small dumb-
bells with very short shafts, and the heads very irregular in shape and covered with
warts. They are "27 — "13 mm. by '21 — '12 mm. The colonies are pale brown colour in
spirit. The spicules are colourless.
Habitat. Straits of Rakaiya, New Britain. Depth, 3 — 4 fathoms.
The figures given by Haeckel in his "Arabische Korallen" 1876, and Klunzinger
(11) of Ammofhea virescens (Sav.) are not by any means alike, but Dr Willey's
specimens closely resemble in form the specimen obtained by Haeckel. The spicules
correspond with the figures given by Klunzinger (Taf II. Fig. 4).
Sub-family II. Siphonogorginae.
12. Chironephthya scoparia, Wright and Studer.
There are only two small fragments of the ends of branches. The larger is
18 mm. long by 2 mm. in diameter and has 10 polyps arranged spirally. The polyp
heads (as in Spongodes) are bent down so that the group of long spicules projects
beyond the head. Each fragment has three polyps near the apex, one being terminal.
vv. IV. 67
502 THE STOLONIFERA AND ALCYONACEA
The tentacular operculum and the collaret agree with Wright and Studer's
description (28). The contracted polyps are I'o mm. in length and l^S mm. in diameter.
The spicules agree in form and size with the description of those of the type
specimen. The branches are yellow and the polyp heads red in colour.
Habitat. Talili Bay, New Britain.
Previously recorded from the Hyalunema ground, 345 fms., Japan. The occurrence
of this species in shallow water off New Britain and in deep water on the Hyalo-
nema ground is a very noteworthy fact in distribution.
Family. Alcyonidae.
Wright and Studer (28, p. 238) are of opinion that only those fleshy Alcyonids
found in the cold and temperate sea should be included in the genus Alcyonium,
those found in the tropical seas being referred to the genus Lohularia.
As these authors have pointed out (28, p. xxi) there is great difficulty in
distinguishing Alcyonium from Savigny's genus Lohularia. It is true that many of
the tropical species have large spicules which cause the cortex to have a leathery
consistency when compared with the cortex of the species of the cold and temperate
seas ; but there appear to us to be many reasons why this feature should not be
regarded as one of generic importance. In this collection there are three specimens
which are so remarkably similar to the European species Alcyonium glomeratum that
it would be absurd to place them in a distinct genus. We agree therefore with
Klunzinger and others in not recognising the genus Lohularia.
A word of explanation may be made here as to the use of the term stalk in
the following pages.
In the genus Sarcophytuin the colony assumes a form which has not inaptly
been compared to that of a mushroom ; there is indeed a well-niarked differentiation
of the upper part or head from the lower supporting part or stalk. The important
morphological difference between the two parts is that the expansible portions of the
polyps, called for the sake of brevity polyp heads, protrude from the capitidum but
do not protrude from the stalk. In some genera of Alcyonidae, such as Alcyonium
and Nephthya, the head and stalk are not so sharply differentiated as in Sarcophytum,
and in the literature of the subject several terras such as " barren part " or " sterile
portion" have been employed for what is morphologically equivalent to the stalk of
Sarcophytum. In our opinion atiy expression which implies barrenness or sterility in
the part named is misleading and consequently we use throughout the term " stalk "
for the part of the colony which does not bear polyp heads.
13. Alcyonium polydactylum, Klunzinger.
The colony has apparently been split in two and only one half is in the bottle.
It is 85 mm. high ; the stem is 55 mm. long by 35 mm. wide, the head is
30 mm. high by 70 mm. broad and is much divided.
There are five main branches, each of which bears several branchlets on wliich
the short, obtuse, somewhat finger-like processes are crowded.
COLLECTED BY DR WILLEY IN NEW BRITAIN, ETC. 503
The main branches are 20 — 40 mm. long and about 20 mm. wide ; the processes
are 5 — 10 mm. long and 3 — 6 mm. wide. The polyps are scattered over the entire
surface of the head at intervals of 1 — 2 mm.
The spicules agree fairly closely with Klunzinger's figures and description. The
long, warted spindles are from o — 2'3 mm. long and "12 — 06 mm. wide. The small
clubs are from '12 — -13 mm. long and 04 — 07 mm. wide.
The colony is fawn colour in spirit.
Habitat. China Straits, British New Guinea.
Previously recorded from the Red Sea (11).
14. Alcyonium glomeratum, Hassall.
There are two complete colonies and one fragment of a species of Alcyonium in
the collection, whk-h appears to be closely related to our British Alcyonium glomer-
atum. They are small, the largest being only 45 mm. in height and 22 mm. wide,
and probably young specimens. Of course we can form no estimate as to the size they
might have grown to when adult.
The larger colony of the two complete ones is pale yellow in colour but here
and there at the polyp heads there may be seen red spots, due probably to clusters
of red spicules. The smaller colony is bright orange-red in colour. The fragment is
intermediate in colour between the two.
In this variation in colour the specimens agree with the European varieties of
A. glomeTatum.
The spicules are of the same general character as those of the European forms.
In the coenenchyma there are some spicules found which are dumb-bell in shape
and the long spindles are larger and apparently more numerous than in the British
form. The Talili spicules are 0-4 mm., the British 0-2 mm., in length. A statement
(10) has been made that in Alcyonium glomeratum dumb-bell shaped spicules are
not found. If this statement ttirned out to be true for all European specimens it
would not in our opinion be sufficient ground for the constitution of a new species
for the Talili specimens which undoubtedly possess them. The spicules of a common
species such as Alcyonium digitatum may vary immensely in size, shape and colour
according to the depth and locality from which they are taken, ami unless we have
a long series of specimens to iudge from, it is unwise to .speak confidently about
the characters of the spicules.
The facts that the spicules are of the same general form as those ot the
European specimens, show similar variations in colour, that the lobes of the colony
are pointed and deeply divided, and that the polyps are of the same size and (in
the retracted condition) of the same appearance, are sufficient when taken together to
justify this determination.
Habitat. Talili Bay, New Britain.
Previously recorded from coasts of Great Britain and Norway.
15. Alcyonium pachyclados, Klunzinger.
There are two small, complete colonics of this form, the larger is 50 mm. by
35 mm. across the capitulum and 15 ram. high. The general appearance, the arrange-
G7— 2
504 THE STOLONIFERA AND ALCYONACEA
ment, shape and size of the " lappets " correspond with those of Klunzinger's specimens.
The spicules agree closely in size and shape.
The colour in spirit is grey.
Habitat. Blanche Bay, New Britain.
Previously recorded from the Red Sea (11).
16. Alcyonium macropodium, n. sp. (PI. L. Figs. 8, 9, 10).
There is one colony of this form. It exhibits a pronounced stalk which is 40 mm.
high by 20mm. and 15mm. in thickness; the capitulum consists of numerous short,
pointed lobes bearing a few large polyps ; it is 4.5 mm. in length and 20 mm. in
diameter. The lobes are about 5 mm. long.
The spicules are warted spindles with a tendency to branch at one end. There
are a few small clubs and four-rayed forms. The spindles vary in size ; they are
2-12 X -368 mm., 1-75 x -24 mm., -736 x -184 mm., -257 x 073 mm., "147 x -0.35 mm. The
four-rayed forms are "163 x '147 mm.
Habitat. Blanche Bay, New Britain.
This species differs very markedly from others of the genus io the long thick stalk
and short branches. These characters with the form and size of the spicules suggest
that it must belong to a distinct species.
17. Sarcophytum ehrenbergi, Marenzeller.
There is only a fragment of the capitulum 70 mm. long by 35 mm. broad. It
is about 10 mm. thick, fairly soft ; the edges bend down somewhat over the stalk in
the form of two small folds. The autozooids are not very close to one another
(7 — 10 in 1 cm.); the pores of the siphonozooids are very minute.
The spicules agree closely in form and size with those described and figured by
Marenzeller (IS).
The colour in spirit is a dirty grey.
Habitat. China Straits, New Guinea. Previously recorded from the Red Sea,
Port Denison (Australia), Viti Islands.
18. Sarcophytum fungiforme, Schenk.
There is one small colony of this form, complete.
The stem is 20 mm. high , and 15 mm. in diameter, the capitulum is 33 mm.
by 25 mm. and 5 mm. thick. There are two deep folds on one side meeting in the
centre of the capitulum. On the other side the capitulum bends down somewhat
over the stalk. The folds are 20 mm. high.
The position and number of the autozooids and siphonozooids per cm. agree with
Schenk's description (24).
The spicules resemble closely in form and size those described and figured by
Schenk. The colour of the colony in spirit is greyish.
Habitat. Blanche Bay, New Britain.
Previously recorded from Ternate.
COLLECTED BY DR WILLEY IN NEW BRITAIN, ETC. 505
19. Sarcophytum glaucuvi, Marenzeller.
There is one complete colony; the stalk is 40 mm. long and 15 mm. in diameter;
the capitulum is 4.5 mm. by 45 mm. and 3 — 5 mm. thick. The whole is decidedly
soft and the capitulum is flabby.
The arrangement of the folds, the distribution of the autozooids and the form
and size of the spicules agree closely with Marenzeller's description (18).
Colour in spirit is brown, the polyps fawn.
Habitat. Blanche Bay, New Britain.
Previously recorded from Tonga Islands and Port Denison and Port Bowen (Australia).
20. Lohophytinn paucifloruin, Ehrenberg.
There are two fairly large specimens, but the basal attachments are missing.
The larger is 50 mm. in height; the stalk is 30 mm. high and 70 mm. broad; the
head is 105 mm. in diameter. It is a somewhat darker brown than the smaller
colony.
Habitat. Talili Bay, New Britain.
Previously recorded from the Red Sea, Amboina and New Zealand (5, 18).
21. Lobophytum marenzelleri, Wright and Studer.
There are two portions of colonies; the larger is 75 mm. in width; the basal
part is missing.
The whole is very firm and rigid.
The capitulum corresponds closely with the description in the Challenger report
(28) as regards the lobation ; it is very hard and packed with very large spicules.
The surface is rough on drying.
The autozooids are scattered over the lobes and where retracted are visible only
as small pores.
The siphonozooids are difficult to see even with a lens. The colour is pale
fawn. The spicules are of about the same length as in the type specimen but are
much narrower. The proportions are : 408 mm. long by '552 mm. wide ; 285 mm.
X -46 mm. ; 1-65 mm. x -33 mm.; -730 mm. x -165 mm. ; -514 mm. x '035 mm.
The small branched clubs are '147 mm. by '0(34 mm.
Habitat. Near Cape Gazelle, New Britain.
Previously recorded from Api, New Hebrides. 60 — 70 fms. (28).
22. Lobophytum densum, Whitelegge.
There is one colony, the basal part of which is missing. It is 70 mm. by 45 mm.
across the head and 40 mm. high.
The lobes are 525 mm. high, by 4 — 8 mm. in their narrow and 7 — 15 mm. in
their broad diameter. The autozooids are about 1 mm. apart. The siphonozooids are
difficult to find even with a lens.
The colony is hard owing to the large number of spicules present. The tuber-
culated spindles are from -8—2 1 mm. long by 17— -31 mm. wide. The .spiny spindles
are from M— 2-2 mm. long by -12— -20 mm. wide. The small clubs are "13 x '03 mm.
506 THE STOLONIFEBA AND ALCYONACEA
The colour is brown with rather darker furrows and pits.
Habitat. Sandal Bay, Lifu ; and China Straits, British New Guinea. Previously
recorded from Funafuti, Ellice Islands (30).
23. Lobophytum crasstmi, Marenzeller.
There is only a fragment of the capitulum of this species measuring 30 mm. by
25 mm. It shows four thick folds. The spicules of the cortex are somewhat larger
than those of the type specimen (18).
The spicules of the coenenchyma are slightly smaller ; they are "23 — '29 mm. long
and '07 — '09 broad. But they are of the same type as those figured by Marenzeller.
Habitat. Loyalty Islands.
Previously recorded from Port Denison (Australia).
ADDENDUM.
Since the above paper was sent to the press my attention has been called to
a memoir by Walther May entitled " Beitrage zur Systematik und Chorologie der
Alcyonaceen" in the Jen. Zeits. f. Naturwiss. xxvi. 1. None of the species described
by us as new agree with the new species described by May. There is no reason to
believe therefore that there is any serious overlapping.
I may point out however that Glavularia flava cannot be accepted as the name
for the new sjaecies described on his page 43, as I employed this name for a species
from Australia described in the Trans. Zool. Soc. Vol. xni. 1895.
S. J. HICKSON.
Feb. 21, 1900.
LITERATURE REFERRED TO.
L AsHwoRTH, J. H. "The structure of Xenia hicksoni." Q. J. M. S., August, 1899.
2. Dana, J. D. "United States Exploring E.xpedition." Zoophytes, 1848.
3. Danielssen, D. C. "Norske Nordhavns E.x^ped. Zool. Alcyonida." 1887.
i. DucHASSAtNG, P. et MicHELOTTi, G. " Memoire sui- les Coralliaires des Antilles." Turin,
1860.
5. Ehrenberg, C. G. "Die Corallthiere des Rothen Meeres." 1831.
6. Gray, J. E. " Some new Alcyonoid Corals in the British Museum." Annals and Maga-
zine of Natural History, Ser. 4, Vol. ill., 1869.
7. HiCKSON, S. J. "A naturalist in North Celebes," 1889, p. 129.
8. HiCKSON, S. J. "The structure and relations of Tubipora." Q. J. M. S., Oct. 1883.
9. HiCKSON, S. J. "Monograph of the Alcyonaria Stolonifera." Trans. Zool. Soc, 1894.
10. HiCKSON, S. J. "The structure of Alcyonium digitatum." Q. J. M. S., 1895.
11. Klunzinger, C. D. "Die Korallthiere des Rothen Meeres." Berlin, 1877.
12. VON Koch, G. "Anatomie des Orgelkoralle." Jena, 1874.
13. VON Koch, G. "Anatomie der Clavularia prolifera, n. sp." Morphologisches Jahrbuch,
vii., 1882.
\
J'
^
.--3
^ ^'» "♦
J
t;N-
S^
J^ ^
.-f
f
tT-
HICKSON AND HILiES. ALCYONACEA.
COLLECTED BY DR WILLEY IN NEW BRITAIN', ETC. 507
14. KoLLiKER, A. "Icones Histiologicae." Leipzig, 1865.
15. KoLLiKER, A. " Die Pennatulide Umbellula und zwei neue typen der Alcyonarien."
Festschrift phys. med. Gesell. Wiirzburg, 1875.
16. KoREN, J. og Danielssex, D. C. "Nye Alcyonider, Goigonider og Pennatulider tilh^rende
Norges Fauna." Bergen, 1883.
17. KiJKENTHAL, W. " Alcyonaceen von Ternate. Nephthyidae und Siphonogorgidae. Abhand-
lungen der Senckenbergischen Naturforschenden Gesellschaft." Frankfurt, Band .xxiii.,
Heft 1, 1896.
18. Marenzeller, E. von. "Ueber die Sarcophytum benannten Alcyoniiden." Zoologische
Jahrbiicher, i., Jena, 1886.
19. MiJLLER, F. "Archiv fur Naturgeschiehte." Jahrgang xxxiii., Bd. 1, 1867.
20. Quoy et Gaimakd. "Voyage de I'Astrolabe." Zoophytes, T. iv. 1833.
21. Ridley, S. O. "Contributions to the knowledge of the Alcyonaria." Annals and
Magazine of Natural History, 1882.
22. Ridley, S. O. "Zoological Collection of H.M.S. 'Alert'." Part I. Melanesian Col-
lections, 1884.
23. Ridley, S. O. " The Alcyonaria of the Mergui Archipelago." Linnaean Society's Journal.
Zoology, Vol. XXI., 1887.
24. Schenk, a. " Clavulariiden, Xeniiden, und Alcyoniiden von Ternate." Abhandl. des
Senckenbergischen Naturforschenden gesellschaft. Band xxiii., Heft 1, 1896.
25. Studer, Th. " On some new species of the genus Spongodes, from the Philippine Islands
and Japanese Seas." Annals and Magazine of Nat. History, Ser. 6, Vol. I., 1888.
26. Studer, Th. Challenger Report on the "Alcyonaria." Supplement, Vol. xxxii., 1889.
27. Studer, Th. " Alc}'onarien aus der Sammlung des Museums in Lubeck." Mitteilungen
des geographischen gesellschaft und des naturhistorischen Museums in Lubeck, II Sec.
Heft 728, 1894.
28. Wright, E. P. and Studer, Th. Cliallenger Report on the " Alcyonaria," Vol. xxxi.
1889.
29. Verrill, a. E. "Report on the Anthozoa and on some additional species dredged by
the 'Blake,' 1877 — 79, and by the U. S. Fish Commission Steamer 'Fish Hawk,'
1880 — 82." Bulletin of the Museum of Comparative Zoology, Harvard, Vol. xi..
No. 1, 1883.
30. Whitelegge, Th. " The Alcyonaria of Funafuti." Memoirs of the Australian ^Museum,
Part 3, 1897.
EXPLANATION OF PLATES L. AND LI.
Fig. 1. Telesto rupicola. Colony x 2. The upper portion is covered by a commensal
sponge througii which the expanded polyps project.
Fig. 2. Spicules of Telesto rupicola.
Fig. 3. Telesto arthuri, n. sp. Colony x 2.
Fig. 4. Spicules of Telesto arthuri. x IG. a. end of .spicule showing size of warts, x 112.
Fig. 5. Spongodes semperi. Colony x 2.
Fig. 6. Spongodes semperi. Tip of branch, showing arrangement of spicules.
Fig. 7. Spicules of Spongodes semjieri.
508 THE STOLOXIFERA AND ALCYONACEA IN NEW BRITAIN, ETC.
Fig. 8. Alcyonium macropodium, n. sp. Colony x 1.
Fig. 9. Alcyoiiium macropodium. One polyp contracted showing valve-like folds (v.) which
can be drawn over the folded tentacles.
Fig. 10. Spicules of Alcyonium macropodium.
PLATE LI.
Fig. 11. Polyp head of Spongodes rakaiyae, n. sp., to show the arrangement of the
spicules.
Fig. 12. A diagrammatic transverse section of a branch of Telesto arthiiri. In the centre
are four large coelenteric cavities {p. c), the tubes of polyps borne close to the part of the
branch from which the section is taken.
The four smaller coelenteric cavities somewhat to the outside of these are polyp tubes
cut nearer to their point of origin ; the polyp tubes become smaller as they are traced
down the branch. The dorsal mesenteries {d. m. f.) with their filaments extend the whole
length of the polyp tubes, and traces of some of the lateral and ventral mesenteries are
usually seen in section. The dorsal mesenteries are on the side of the tube nearest to the
axis.
Ova {ov.) are found attaclied to some of the lateral mesenteries.
The mesogloea between the polyp tubes contains numerous small endoderm-canals, but
these do not extend to the surface of the branch (c. c).
Two pohps are shown cut across. One {t. s.) is a transverse section of the polyp just
below the stomodaeum, sliowing the large bundles of muscle fibres on the mesenteries which
however extend only for a very short distance down the tube, and the mesenteries become
mere little projecting lumps of mesogloea covered by a thin layer of endoderm cells con-
tinuous with the layer lining the polyp tube.
The other polyp (/. *■.) is cut longitudinally and shows two retracted tentacles and the
stomodaeum in longitudinal section.
The large cavities near the edge of the section represent the ca^-ities occupied in nature
by the spicules.
Fig. 13. A retracted polyp of Clavularia viridis as seen when cut longitudinally in
half. V. m. /., the ventro-lateral mesenterial filaments commencing some distance from the
stomodaeum ; d. m. f., the dorsal mesenterial filaments extending the whole distance from the
stomodaeum to the base; </., the gonads; t., tentacle; St., Stomodaeum.
Fig. 14. A portion of a dorsal mesentery of a polyp of Clavularia viridis enlarged to
show the form of the dorsal mesenterial filament [d. m. /.).
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WflBt.NewrriiiT;
HICI^ON AND HILiES. ALCYOITACEA.
REPORT ON THE XENIIDAE COLLECTED BY DR WILLEY.
By J. H. ASHWORTH, D.Sc,
Demonstrator in Zoology, Owens College, Manchester.
With Plates LII. and LIII.
CONTENTS.
PAGE
Introduction 509
Xenia crassa, Schenk ................ 510
X. memhraaacea, Schenk ............... 512
X. umhellata, Savigny 513
X. viridis, Schenk 516
New Species. X. Novae-Britanniae, Ash worth 518
Diagnosis of the Species Xenia Xovae-Britanniae, Ashworth ........ 521
Habits, Distribution and Classification of the Xeiiiidae ......... 522
General Internal Anatomy ............... 523
Stomodaeum and Mesenterial Filaments ............ 523
Coelentera of Polyps 525
Mesogloea, its Canals and Cells 525
Ectoderm 526
Endoderm .................. 526
Gonads 526
Literature 528
Dr Willey's collection of specimens of the genus Xenia contains fifteen colonies
which may be referred to five species.
Of the sixteen species of Xenia hitherto described eight were founded by
Schenk (1S96)' upon specimens brought from Ternate (Moluccas). Three of the
specimens in Dr Willey's collection belong to three of these newly described species,
which are now recorded for the second time only. One of the other two species is
moderately well-known, being, in fact, the fir.st described species of Xenia (X. unibellaia,
Savigny) ; the other species is new and is represented in the collection by eight
specimens. Thus although four of the species have been previously described three of
them have hitherto been met with only once, and as all the specimens are of considerable
interest a brief description of each is given below.
' The dates in brackets form references to the list of papers given on p. 528.
W. IV. 68
510 KEPORT OX THE XEXIIDAE COLLECTED BY DR "WILLEY.
The specimens referred to the three species X. umbellata, X. crassa, and X.
membranacea differ in one or more characters from the specimens described as tj-pical
of those species, but the differences are certainly not of sufficient importance to
justif)' the formation of new species. For example, the differences between the
specimen which is referred to the species X. crassa and the type specimen described
by Schenk, may be accounted for by the fact that Dr Willey's specimen is a young
colony the polyps of which have not yet acquired their adult size and characters,
while Schenk's was probably a more mature colony. It is therefore important to examine
and record the condition of the genital products of a colony in order that it may be
ascertained whether the polyps are, or are not, adult. Some of the differences between
Klunzinger's specimen of X. umbellata and the colony of this species in Dr Willey's
collection may also be attributed to differences in age (see p. 515), while others, e.g., the
form of the colony, may be due to the influence of dissimilar habitat upon the mode of
growth. It seems reasonable to suppose that specimens obtained from the Red Sea (eis
was Klunzinger's X. umbellata) and from New Britain may be subjected to rather
different external conditions, which may have an influence upon the colony, producing
variation in its general form, the stoutness of its polyps, etc. The photographs
coutained in Professor Hickson's Report on the specimens of Millepora collected by
Dr Willev show how manv different forms or facies mav be assumed bv a sinsjle
species. Some attention has been devoted to the anatomy of each species, but as
they all agree in their main features with X. Hicksoni, the anatomy of which has
been described in detail (Ashworth, 1S99) only a few notes on their structure are
appended. Important confirmation has been obtained of all the chief features de-
scribed in X. Hicksoni, with the exception of the giant flagella, which do not appear
to be present in any of these species. The most important point confirmed is
certainly the absence of ventral and lateral mesenterial filaments in which the polyps
of Xenia and Heteroxenia differ from those (autozooids) of any other Alcyonarian
hitherto described.
XENIA CRASSA. Schenk.
This species is represented by a small and probably young colony, the single
unbranched stem of which is thick, soft and fleshy and somewhat knob-shaped, its
polyp-bearing summit being convex. The height of the stem is 10 mm. — 11 mm. ;
its diameter at the base is 10 mm. and at the summit 15 mm.
The polyps' are short and, stout and stand close together, especially round the
edge of the summit of the stem. The measurements of the larger polyps are — ■
body of the polyp 3 mm. long and 1-5 mm. — 2 mm. broad (a very few polyps are
2-5 mm. in breadth), tentacles 2-5 mm. — 3-2 mm. long and 7 mm. — 12 mm. wide. Each
' In order that the measurements and characters given below, may be compared with those given by
previous authors, the word polyp is used, in this and in similar connections in this paper, in the same sense
as these authors have used it, viz., to indicate the free or exsert portion of the polyp. It should be remembered
however that this is not the whole of the polyp, a considerable portion of it is enclosed in the stem from
the summit of which the free portion of the polyp projects, indeed a primary polyp (i.e. one of the first
formed polyps of the colony) extends down to the base of the colony. (See Plate LII. Fig. 7.)
REPORT ON THE XENIIDAE COLLECTED BY DR WILLEY. 5 1 1
of the somewhat short and wide tentacles hears on its inner face three rows of pinnules
on each side of the middle line. There are nine to twelve pinnides in each row. In
the majority of the polyps the pinnules occupy the whole of the inner face of the
tentacle with the e.xception of a small area in the middle line near the base. The
pinnules, which are rather stout and conical, are o mm. — '7 mm. in length and '2 mm.
in width.
Besides these well-developed polj'ps there are man}' buds or young polyp.s in
various stages of development. These are situated not only round the outer edge of
the polyp-bearing summit of the stem, where thej' are most usually found in other
Xeniidae, but are also scattered over the convex end of the stem between the bases
of the larger and older polyps. The buds are, however, much more numerous round
the edge of the convex disc, there being about thirty polyps less than one millimetre
in length situated around this edge. In the youngest polyps, the length of which is
"4 mm., the tentacles are indicated by eight small rounded elevations separated from
each other by slight grooves. In slightly older polyps '7 mm. long the tentacles are
simple rounded wart-like elevations "3 mm. high. The first pinnules appear when the
tentacles of the polyp attain a length of about 4 mm., and from this stage onwards
there is a gradual increase in the number of jDiunules along with the increase in length
of the tentacle, e.ff. a tentacle '5 mm. long shows when seen from the outer aspect
three pinnules on each side of the middle line, a tentacle '75 mm. long shows five
pinnules, one '9 mm. long shows six pinnules, &c.
The disc-like spicules are extraordinarily numerous, and in many parts they are in
contact with or even overlapping each other. They are whitish by reflected light and
either light yellow or light reddish-brown by transmitted light. Most of them are oval
in shape but some circular ones are present. They are 018 mm. — 024 mm. in length
and '015 mm. — "018 mm. in breadth.
The stem of the colony is light yellow-brown, slightly tinged with green in the
upper part, the polyps are a much lighter shade of the same colour. The greyish-
white colour of the tentacles and pinnules is probably due in a great measure to
the light reflected from the very numerous spicules present in those parts.
The specimen is a male but the gonads are small. The sperm sacs are recognisable
as .spherical swellings on the edges of the six ventral and lateral mesenteries, but
the largest sacs present are only "08 mm. in diameter, and are obviously in an early
stage of development, the cells which they contain having undergone comparatively
few divisions. These sacs are similar to the largest of the three of X. Hicksoni
represented on PI. 27, Fig. 30 (1899).
Two large polyps were stained and cleared. The stomodaeum of these polyps
is about '9 mm. long, and the wall of each shows a considerable number of small
light areas which probably indicate the presence of swollen empty cells similar to
those which have been described in the stomodaeum of X. Hicksoni (1899, p. 251).
Habitat : — Isle du Phare Reef, Noumea, New Caledonia.
The specimen agrees in most respects with the diagnosis of A', crassa, Schenk (1896,
p. .58), but the dimensions of the polyps of Dr VVilley's specimen are about two-thirds
those of Schenk's specimen and the number of pinnules on the tentacles differs
68—2
512 REPORT ON THE XEXIIDAE COLLECTED BY DR WILLEY.
in nearly the same proportion (9 — 12 in each row in Dr Willey's, 15 — 18 in Schenk's).
These differences are probably due to the fact that this specimen is a young colony
the polyps of which have not yet grown to their full size and the tentacles of which
have not yet acquired the full number of pinnules. (This is supported by the
condition of the genital products described above.) In spite of these differences the
characters of the colony, e.g. the shape and character of the stem, the form and
structure of the polj^s, the shape of the pinnules and the extraordinary abundance
of spicules, are in close agreement with the corresponding characters of Schenk's
species.
This species has been previously recorded from Ternate (Moluccas).
XENIA MEMBRANACEA. Schenk.
This colony consists of two almost equal and parallel stems arising from a common
basal membrane which is attached to the two branches of a V-shaped, piece of dead
madrepore. The membrane which has spread out over the adjacent surface of the
madrepore is thickest around the bases of the stems, becoming gradually thinner towards
its free edge, where it is about a millimetre thick. The stems are 15 mm. long and
10 mm. in diameter, and 18 mm. long and 9 mm. in diameter respectively. The free
end of each stem is slightly convex and bears the polyps, which are numerous and
closely packed together, often being somewhat flattened by mutual pressure.
The polyps are moderately long and have a slender appearance. The body of
each polyp is 5 mm. — 10 mm. in length (a very few reach 12 mm. in length) and
•8 mm. — 12 mm. in breadth. The tentacles are also long, slender and pointed, being
4"5 mm. — 7 mm. in length and about '4 mm. — "8 mm. broad. The inner face of each
tentacle bears rather long and thin pinnules, which are arranged, but not very clearly,
in six rows. The pinnules generally occupy the whole of the inner face of the
tentacle, but in a few polyps the middle line of the tentacle for a short distance
from the base is free from pinnules. There are, on an average, 16 to 18 pinnules
in each row, but in some of the longest tentacles there are 24 in each row. In
the middle of the tentacle, where they are largest, the pinnules are "5 mm. — 6 mm.
in length and about "1 mm. in breadth.
In this specimen also buds are present but the}- are much fewer in number
than in the preceding colony (X. crassa). They are found only on the edge of the
polyp-bearing summit of each stem. The smallest is "4 mm. long and its tentacles
are but faintly indicated. In a, bud '6 mm. long the tentacles are finger-shaped lobes
•25 mm. long, while in a polyp 1 mm. long they have attained a length of So mm.
but none of them yet possess pinnules. A young polyp 1-4 mm. long has advanced
considerably, as its tentacles are '6 mm. long and show two or three pinnules on
each side of the middle line of the tentacle (seen from the outer aspect), while a
young pol\i3 25 mm. long bears tentacles 13 mm. in length, each of which shows on
its outer face 6 — 7 pinnules on each side of the middle line.
The spicules are round or oval discs '012 mm. — -018 mm. in length and 01 mm. —
•012 mm. in breadth. They are either yellowish or slightly bluish-white by reflected
light and many of them show iridescent colours. By transmitted light the spicules
REPORT ON THE XENIIDAE COLLECTED BY DR WILLEY. 513
are yellowish-brown. They are present in moderate numbers in the stem, but are
more numerous in the body of the polyp and in the tentacles, and still more
numerous on the outer face of the pinnules, but there are comparative!}- few on the
inner face of the pinnules.
The specimen is dark brown, its colour being due partly to the spicules and to
the contained zooxanthellae and partly to the colour of the soft tissues.
The stomodaeum of the lai-ger polyps is 1"7 mm. — 1'9 mm. in length.
On examining in spirit the surface of the ectoderm of the polyps, numerous
minute refringent oval bodies are clearly visible. Most of these bodies are imbedded
in, and their long axes are at right angles to the free surface of, the ectoderm.
These are very probably nematocysts. They are '008 mm. — '009 mm. in length and
■002.5 mm. in diameter, thus agreeing very closely with the nematocysts of X. Hicksoni
(1S99, p. 258), which are "008 mm. long and "002 mm. — '003 mm. in diameter.
The specimen is apparently a male, but the sperm sacs are very few in number
and very small in size, and are obviously in an early stage of development. They
are rather oval and the largest measure only about '05 mm. along their greater
diameter.
Habitat. New Britain.
This specimen agrees with the description given by Schenk of X. rnembi-anacea.
The general characters of the stem and polyps agree with the diagnosis of the species
very well indeed ; the polyps and tentacles of Dr Willey's specimen are rather larger
than those of the type specimen, but their general proportions are almost identical.
Schenk's species derived its name from the basal membranous expanse which bound
together the bases of the stems. A similar basal membrane is present in Dr Wille3f's
specimen but is developed to a less extent, probably owing to the dififerent mode of
attachment of the colony. Schenk's specimen was attached to a sandy (and probably flat)
substratum which would offer favourable opportunities for the formation of the basal
membrane.
Previously recorded from Ternate (Moluccas).
XENIA UMBELLATA. Savigny.
There are three colonies referable to this species.
I. The basal portion of the largest colony encrusts a piece of weathered coral
rock. From this rather thick membranous base (which is 2 mm. — 3 mm. in thick-
ness) three stout fleshy stems arise, the largest of which is about 33 mm. high
and 30 mm. x 15 mm. at the base. At a distance of about 10 mm. from the summit
this stem is divided into a larger and a smaller branch by a deep fissure. The stenj
next in point of size is 25 mm. long and measures 15 mm. x 15 mm. at the base.
The smallest of the three stems of this colony is 20 mm. high and 15 mm. x 13 mm. at
the base. All the stems increase slightly in diameter as they ascend. The summit
of each stem is a flat or slightly convex area from which the polyps arise.
II. The second colony is also attached at the base to a piece of worn coral
rock. It consists of a single stout fleshy stem about 25 mm. high and 23 x 13 mm.
514 REPORT ON THE XENIIDAE COLLECTED BY DR WILLEY.
at the base. The stem widens as it ascends and its breadth at the summit is
33 mm. x 16 mm. The pol3rp-bearing area is slightly convex. The base of the colony
is prolonged downwards at the edges, forming an almost hemispherical cap which is
closely applied to the rock on which the colony is fixed.
HI. The smallest colony consists of a single stem about 15 mm. long and about
12 mm. X 10 mm. at the base.
The polyps arise moderately close together on the end of each stem. The bodies
of the fully developed ones are 5 mm. — 10 mm. long (a few reach 13 mm.) and
1"2 mm. — 1-8 mm. wide. The tentacles are long, being 5 mm. — 8 mm. in length and
1"3 mm. — 16 mm. broad, and bear on each side of the middle line three rows of
pinnules of about 2"2 — 29 in each row. The pinnules are long and slender, ".5 mm. —
"8 mm. in length and 1 mm. — '12 mm. in width. The inner face of each tentacle
presents an area free from pinnules. This area is about "4 mm. wide at the base of
the tentacle, gradually narrows towards its tip, and is no longer recognisable in
the distal fourth of the tentacle ; in this region the pinnules of the two sides are in
contact at their bases.
Buds are found not only round the edge of the summit of the stem but also
scattered over the whole of the summit between the bases of the older polyps.
Those round the edge of the summit are similar to the buds found in other species
of Xenia (PI. LIII. Fig. 10), except that the young polyp is l^B mm. — l^S mm. long and
its tentacles have reached a length of nearly 1 mm. before the first pinnules appear
upon them. In the other species of Xenia the pinnules appear when the polyp is
much smaller and its tentacles are only "4 mm. — '6 mm. long. There are on the ends
of two of the large stems three small areas in each of which four or five buds are
found. Several of these buds differ from the normal buds present round the edge
of the summit of the stem, the body of the former being much larger in jH-opor-
tion to the tentacles than is usual. One of these buds is represented in Fig. 12.
Its total length is 5 mm., its stout finger-shaped, rather unequal tentacles are only
1"0 mm. to 1'3 mm. long. Three of them are trilobed at their tips, i.e. there is an
indication of the formation of the first two pinnules, one on each side of the axis
of the tentacle. Another similar bud G nun. long has slender digitiform tentacles
17 mm. long, each of which bears one or two pinnules on each side of the middle
line (see Fig. 11). In these and other similar buds from the three areas mentioned
above, the tentacles are onh- about one-fourth of the total length of the polyp and
bear few pinnules, whereas in - buds of similar size present round the edge of the
summit of the stem the tentacles are about one-half the total length of the polyp and
bear a larger number of pinnules. (See Figs. 10 and 11 ; the tentacles of the two young
polyps rejjresented are equal in length.)
Few spicules are present in the stem except at the base, where they are numerous
and stand almost in contact with each other. The ■pre.sence of many spicules in this
region gives additional strength and rigidity to the basal portion by which the colon}' is
attached to the rock. Spicules are moderately numerous in the body of the polyp,
becoming more numerous towards its distal end. They are abundant on the outer
faces of the tentacles and pinnules, being close together in the latter. The\' are less
REPORT ON THE XENIIDAE COLLECTED BY DR WILLEY. 515
numerous on the inner faces of the tentacles and pinnules. By reflected light the
spicules are whitish or very slightly yellow. By transmitted light some of the spicules
are almost colourless but the majority have a moderately strong reddish-brown colour.
The spicules are oval discs OIQ mm. — '018 mm. in length and '01 mm. — 014 mm. broad.
The greater part of the colony is yellow in colour, but those parts which have
been closely pressed together and partly protected from the action of the spirit are
reddish-brown in coloui-. The label on the bottle indicates that this brown colour
predominated in life.
Ova are present in considerable numbers on the edges of the ventral and lateral
mesenteries of many of the larger coelentera. The largest oya, which are somewhat
oval in shape, are 'o mm. x "6 mm. in diameter (Fig. 13). These are probably mature
ova. The spherical or oval nucleus is Oo mm. to '07 mm. in diameter and tlie exceedingly
fine and close network which it contains stains deeply with haemato.xylin, safranin, &c.
In each nucleus there are about 15 to 20 spherical very deeply staining bodies about
iifi in diameter, and a larger number of similar but much smaller bodies about Ifj. in
diameter. The nucleus is situated near the edge or circumference of the ovum. A
germinal spot is not present in large ova but is very well marked in young ova
until they attain a diameter of about "12 mm. In ova of this size and in younger
ones the germinal vesicle and germinal spot are quite typical, but in ova gi-eater
than '12 mm. in diameter the germinal spot is either very indistinct or quite absent
and the nucleus has assumed the appearance and structure of the nucleus of the
ripe ovum described above. The protoplasm of the peripheral part of the ovum is
finely granular and devoid of yolk granules, which are however present in large
quantity in the central part of the ovum. In sections of preserved ova, the yolk
substance of which has been dissolved out, the protoplasm of the inner portion
contains numerous more or less spherical cavities which in life contained the yolk
spherules. This portion of the ovum presents a reticulate appearance as shown in the
figure (Fig. 13), the protoplasmic strands are granular but the granules are not ver}'
obvious except in the peripheral portion of the inner or yolk-containmg protoplasm,
where there are several small but deeply staining granules. The surface of the ovum
is depressed in the region of the germinal vesicle. These ova are rather larger than
those of Alcyonium digitatum but in other respects resemble them closely (cf Hickson,
1895, p. 377, and Figs. 40, 41).
The stomodaeum of the fully developed polyps is 1'7 mm. to 2-0 mm. in length.
Habitat. Found at a depth of two or three fathoms in Blanche Bay. New Britain.
On first examining these colonies they appeared to belong to none of the hitherto
described species of Xenia. On comparing their chief characters with those of other
species it was found that they were most nearly allied to X. umbellata, Savigny.
The general appearance of the colony, the proportionate measurements and shape
of the parts of the polyps, the long slender pinnules — all agree with Klunzinger's
description of X. umhellata (1877, p. 39). There are however some differences which
are mentioned and considered below —
(1) The polyps are slightly larger than those of Klunzinger's specimen, but
the proportion of length to width is practically identical in both.
516 REPORT ON THE XEXIIDAE COLLECTED BY DR WILLEY.
(2) The tentacles of these specimens are 5 mm. — 8 mm. long, while those of
Klunzinger's specimen are only 3 mm. — 5 mm. in length, but Dana states that " in
Savigny's figure the tentacles are a third of an inch long " (1848, p. 604), i.e. 8 mm.
The tentacles of Dr Willey's specimens agree very closely with the latter diagnosis.
(3) There is a larger number (22 — 29) of pinnules in these specimens than in
Klunzinger's specimen (12 — 15), but this is readily explained when the greater length
of the tentacles of the former is considered. As shown in X. Hicksoni (1899, pp. 283,
284) there is a gradual and moderately constant proportional increase in the number
of pinnules as the tentacles grow in length, and the proportion which holds between
the length of the tentacles and the number of pinnules is almost identical in the
two s])ecimens under consideration. As shown above the larger polyps of Dr Willey's
specimens contain ripe ova, and therefore the polyps are sexually mature and have
probably reached their full size. There is no record of the maturity or other\vise
of Klunzinger's specimen, and it is possible that the poh-ps which he measured had
not attained their full size, which would account for the differences between his specimen
and those of Savigny in the size of the tentacles and in the number of their pinnules.
4. Klunzinger's figure (1877, Tafel III. fig. 3 a) which shows a broad area free
from pinnules on the inner face of the tentacle does not agree with other diagnoses
of the species X umhellata, e.g. cf Dana, p. 605. " The papillae in the upper view
of the tentacle are separated by an extremely narrow line so that those of the two
sides almost meet at the base." The tentacles of the specimens from New Britain agree
with the latter description very closely.
5. The stems are thicker and the whole colony has a rather stouter appear-
ance than most specimens of X. umhellata, but this may be due to the influence of
difi"erent external conditions and it is certainly not a sufficient reason for separating
this species from X. umhellata.
It may therefore be concluded that although the specimens differ slightly from
Klunzinger's description there is no essential point of difference between them and
X. umhellata. The variations may all be accounted for by differences in age and habitat.
Previously recorded from the Red Sea, where it is abundant in the shallow waters of
the coral reefs, growing on rocks or old coral branches.
XEXIA VI EI BIS. ScHESK.
This species is represented in the collection by a portion of a colony, which consists
of a single unbranched stem nearly half of which has been cut or torn away longi-
tudinally. The stem is very thick, soft and fleshy, and of nearly uniform diameter
along its whole length. It is 18 mm. long and 22 mm. wide at the free end. The
surface of the stem is almost smooth, but near the distal end slight longitudinal ridges
mark the position of the outermost or peripheral coelentera of the polyps. The summit
of the stem is almost flat or only slightly convex.
The poh'ps are short and stout and moderately numerous. They are 4 mm. — 6 mm.
long and 1'5 mm. — 2 mm. broad, (a very stout polyp measures 2'5 mm. in diameter).
The tentacles also are short and thick, they are 4 mm. — 6 mm. in length and 1 mm. —
REPORT ON THE XENIIDAE COLLECTED BY DR WILLEY. 517
I'o mm. in breadth. The pinnules on the tentacle are beautifully and regularly arranged
in three rows on each side of the middle line of the tentacle. Each row contains
15 — 20 pinnules. Those near the base of the tentacle, the first four or six transverse
rows, are short rounded elevations or "warts" (Schenk) about "2 mm. — 3 mm. in length
and "15 mm. — '2 ram. in breadth, but the more distal ones are conical, thick, somewhat
pointed pinnules 5 mm. — 7 mm. long and "2 mm. — •25 mm. in diameter at their thickest
part. The middle line of the inner face of the tentacle is free from pinnules in the
proximal three-fourths of its length. The outer face of the tentacle is markedly convex.
Round the edge of the polyp-bearing summit of the stem and also on other portions
of the summit between the bases of the larger polyps, young polyps in various stages of
development may be found. They are more numerous on the edge of the summit. Most
of them already show pinnules upon their tentacles, but those polyps the total length of
which does not exceed IS mm. (of which the tentacles form about -5 mm.) do not yet
show pinnules upon their tentacles. "
The spicules are disc-shaped and numerous. They are generally oval in shape,
•018 mm. — ■02 mm. in length, about -Ol-smm. in breadth and OOC mm. in thickness. They
are whitish by reflected light and yellowish or yellowish-brown by transmitted light.
Spicules are moderately numerous in the stem, there are rather more in the body of the
polyp and still more on the outer face of the tentacles and pinnules. Spicules are less
numerous on the inner than on the outer face of the tentacles and pinnules.
The stem is greyish green in colour and the polyps a much lighter shade of the
same colour. The tentacles are almost -white but have the slightest tinge of brown.
Sperm sacs are present though not in very large numbers on the edges of the
ventral and lateral mesenteries of the larger polyps. The youngest sperm sacs form
small spherical projections at the side or free edge of the mesentery, each of which
consists of a follicle of endoderm cells within which is a thin lamina of mesogloea
enclosing the genital cell or the cells which have been produced by its division (see
PI. LIII. Fig. 14). The largest sperm sacs are -25 mm. in diameter and the spermatozoa
which they contain, though in an advanced stage of development, are not ripe.
On carefully examining the mesenteries in sections, many of them are seen to bear
young ova, each of which is enclosed in an oval or pear-shaped follicle, in addition
to the sperm sacs. (See Fig. 14.) Each ovum has rather deeply staining, somewhat
granular protoplasm, scattered through which are several small cavities which, in life,
probably contained the yolk gi-anules of the ovum. The nucleus is large, clear and
vesicular, about •OI.t mm. in diameter, and contains a well-marked, spherical, deeply
staining nucleolus. These young ova are generally from 03 mm. to 'O-S mm. in diameter
but one or two attain a diameter of "08 mm. This colony is therefore hermaphrodite, but
it is evident that the male products will ripen first as they are in a much more advanced
stage of develo])ment than the ova.
The stomodaeum of the larger polyps is 21 nun.— 2^2 mm. long. The goblet-like
cells to which reference has been made above may be seen among the ordinary
columnar cells in the ventral and lateral regions of the stomodaeum, especially near its
inner end. The siphonogl)-ph extends along the inner third (i.e. '7 nun. — •S mm.) of its
length.
w. IV. «9
518 REPORT ON THE XENIIDAE COLLECTED BY DR WILLEY.
Habitat. Isle du Phare Reef, Noumea, New Caledonia.
This specimen certainly belongs to the species X. viridis, Schenk. Even the more
unimportant diagnostic characters of Schenk's species may be recognised in this specimen,
e.g. the strongly convex outer face of the tentacle, the thick soft stem, rich in coenenchym,
and the cup shape of the distal end of some of the polyps.
Previously recorded from Ternate (Moluccas).
NEW SPECIES.
XENIA NOVAE BRITAXNIAE, sp. nov.
There are eight specimens referable to this species, all of which are very similar in
shape, being dome-like or knob-shaped. The stem of the colony is usually single and
unbranched, but each of the two colonies from Lifu, marked ill. and D II. (see table
of measurements on p. .519), consists of two almost equal stems connected at their bases
by a thin flat band or membrane. The stem of the largest colony is 18 mm. high,
2.5 mm. x 15 mm. at the base and 30 mm. x 25 mm. at the summit. The .smallest colony
is about 5 mm. high and 8 mm. broad.
The pol}'ps of all the specimens are small and resemble each other very closely in
the measurements of their various parts. The measurements of the fully-developed polyps
are: — body of the polyp 2-8 mm.- — 4o mm. long (a very few reach 5 mm. in length) and
•8 mm. — r3 mm. broad; tentacles 18 mm. — 3'5 mm. long and "7 mm — '8 nmi. (occasionally
1"0 mm.) broad. Each tentacle beai's three rows of short pinnules on each side of the
middle line. There are 8 — 12 pinnules in each row. The pinnules at the proximal
end of the tentacle are much shorter than the more distal ones. Seen from the
outer aspect the proximal three or four pinnules are small wart-like protuberances
■17 mm. — "2 mm. long and '12 mm. — '18 mm. in diameter, while the moi-e distal ones
are longer and more typical pinnules "2 nmi. — So mm. in length and '15 mm. — "2 mm.
in diameter. They are largest in the middle of the tentacle and decrease slightly in
size towards its tip (see Plate LII. Fig. 2). Seen from the inner aspect the pinnules are
wart-like or club-shaped outgrowths standing almost at right angles to the face of the
tentacle (see Fig. 3). On the inner face of the proximal half of the tentacle there is
in the middle line a narrow area free from pinnules, but nearer the tip of the tentacle
the pinnules stand quite close together and many of them are flattened, on one or more
of their faces, by mutual pressure.
In all the specimens buds occur apparently only on the edge of the summit of
the stem. These buds are similar throughout all the colonies. The smallest bud found
is "6 mm. long and its tentacles are simple rounded lobes '25 mm. long. A specimen
1-1 mm. long has simple but rather tiuger-shaped tentacles '4 mm. long. In a young
polyp 1^2 mm. long each of the tentacles is ••5 mm. in length and bears a small pinnule
on each side of the middle line. In a rather older polyp 1-7 mm. in length each of
the tentacles is '8 mm. long and bears two pinnules on each side.
Spicules are abundant. They are generally oval discs measuring "2 mm. — 22 mm.
along their greater diameter, '015 mm. — -018 mm. along their smaller diameter and
KEPORT ON THE XENIIDAE COLLECTED BY DR WILLEY.
519
•004 mm. in thickness. They are white or slightly bluish-white by reflected light but
light reddish-brown by transmitted light. There are comparatively few spicules in the
stem but they are more numerous in the body of the polyp, especially towards its
distal end around the bases of the tentacles. They are abundant in the tentacles
and very numerous on the outer face of the jDinnules, where they are almost in
contact with each other. (See Figs. 5 and 6.)
All the stems of the specimens are a very pale yellowish -green colour, but the
polyps are whitish with a pale bluish bloom, due to the very numerous spicules which
they contain.
Three of the colonies have been examined for sexual products and all proved to
be males. On the mesenteries of many of the larger polyps sperm sacs are present
in considerable numbers, in some cases they are so numerous that they almost fill
up the cavity of the coelenteron in which they are contained. The largest sperm sacs
are about -32 mm. in diameter and contain almost ripe spermatozoa. (The ripe sperm
sacs of X. Hicksoni measure -35 mm. in diameter.) The sperm sacs are found only in
those portions of the coelentera contained in the upper o mm. of the stem. (See Fig. 7.)
From each of the colonies two or three of the largest polyps were removed, stained and
cleared. The stomodaeum is very uniform in length throughout. Its length averages as
nearly as pqssible 1 mm. In some of the smaller polyps (the body of which is only
about 3 mm. long) the stomodaeum is "8 mm. long, while in the largest polyps (the body
of which is about 5 mm. long) it is about 1-2 mm. in length. In one of the most
favourable specimens, cells very similar to the goblet cells to which reference has been
made above may be seen in the ventral and lateral walls of the stomodaeum.
Appended are the details of the various colonies : — •
Colony
Height of
stem of
colony
Measurements
at base
of stem
Measurements
at summit
of stem
Talili Bay I.
mm.
16
mm
20x11
mm.
25x11
II.
9
10x10
12x9
„ III.
5
8x5
8x6
Lifu A.
18
20x17
25x15
„ B.
18
25x15
30x25
„ C.
25
lGxl.3
22x20
„ D. I.
22—23
17-18
17-18
„ D. II.
8-9
18x13
18x13
The colonics from Talili Bay have a very convex polyp-bearing surface. Their
polyps are slightly smaller than those of the specimens from Lifu, the bodies of the
69—2
520 REPORT ON THE XEXIIDAE COLLECTED BY DR WILLEY.
fully developed ones being 2'8 mm. — 3'3 mm. long and 'Smm. — I'l mm. in diameter,
and their tentacles IS mm. — 2'1 mm. long and about '8 mm. wide.
The colonies from Lifu are all very similar, being knob-like or dome-shaped
colonies, though one or two, e.g. B ami D ii. are rather flattened from above down-
wards, and hence the polyp-bearing summit of the stem in these examples is rather
less conve.K than in the others. The fully-developed polyps of all these are moderately
uniform in size and character. Their measurements are: — body of polyp 3 mm.— .5 mm.
long and '8 mm. — 1'3 mm. broad, tentacles 2-2 mm. — 3'o mm. long and '7 mm. — 8 mm.
(occasionally I'O mm.) wide.
The colony from Lifu marked D i. consists of two .similar and almost equal
dome-shaped stems, the measurements of which are given in the above table (p. 519).
The stems are bound together at their bases by a thin membrane.
Two rather irregular and flattened stems joined together at their bases by a
slender connection form the colony marked D il. In this colony also the two stems are
almost equal and the measurements given in the table apply to both.
The largest polyps of this species are found on the colony marked D i. Several
of these are 8 mm. in length (body .5 mm., tentacles 3 mm.).
These specimens do not agree with the descriptions of any of the known species
of Xenia. They differ from most other species in their smaller polyps and short
rounded pinnules, and the latter are also much fewer in number than in most of
the known species. In some respects these specimens resemble X. plicata, Schenk,
and X. Garciae, Bourne, but on careful examination several important distinctions are
recognised. Dr Willey's specimens differ from X. plicata, in —
(1) Their smaller polyps, the body of which is 2-8 mm. — 4'5 mm. long and
"8 mm. — rS mm. broad (in A', plicata the corresponding measurements are 4 mm. —
5 mm. and l'-5 mm. — 2 mm.).
(2) Their much shorter tentacles, which are onh' 1"S mm. — 3 5 mm. long, while
those of X. plicata are 5 mm. — 7 mm. long.
(3) The smaller number of pinnules on the tentacles, there being only 8 — 12
in each row while there are 18 — 22 in each row in X. plicata.
There are other differences, e.g. in the arrangement of the polyps on the summit
of the colony, &c., but the above are the chief
After comparing Dr WilleyV specimens with Mr Bourne's figure and description of
X. Garciae, I was still in doubt whether the two might not be identical. Mr
Bourne has kindly lent to me his specimen of X. Garciae so that I have been able
to compare the two directly. Dr Willey's specimens certainly do not belong to this
species. They differ from it in —
(Ij Their longer polyps, the body of which is 28 mm. — 45 mm. long (the
corresponding measurement in X. Garciae is 1"8 mm. — 3 mm.).
(2) Their longer tentacles, which attain a length of 1"8 mm. — ^o"5mm. while those
of X. Garciae are not more than 2 mm. in length.
EEPORT ON THE XENIIDAE COLLECTED BY DR WILLEY. 521
(3) Their shorter pinnules, the length of which is '2 mm. — 'So mm. compared with
•35 mm. — '5 mm. in X. Garciae.
The pinnules of these two forms also differ in their shape and in their position
on the tentacle. The pinnules of Dr Willey's specimens are not only shorter than
those of X. Garciae but they are quite rounded at their tips, while those of the latter
are almost pointed. On viewing the outer face of the tentacles of X. Garciae it is
at once noticed that the pinnules which are then visible arise close together near the
middle line (see Fig. 4 and 1896, Plate 12, Fig. 14). In this respect X. Garciae
differs markedly from Dr Willey's specimens and indeed from any other species of Xenia
hitherto described (except Bourne's specimen of A', coerulea, in which also the long
slender pinnules on the outer face of the tentacles arise moderately close to the
middle line).
A comparison has been made above between the specimens from this collection
and the only two hitherto described .species to which they may be said to have any
resemblance and it is evident that although there may be certain superficial resem-
blances between Dr Willey's specimens and X. plicata and X. Garciae the former differ
in several important characters from these and from any other hitherto described species.
It cannot be argued that these are young colonies, the polyps of which have
not yet attained their adult length and the tentacles of which have not yet acquired
their full number of pinnules, because even the colony from Talili Bay, the largest
polyps of which are considerably smaller than those of the colonies from Lifu, con-
tains sperm sacs which are almost, if not quite, ripe. (The diameter of these sacs is
■28 mm., the diameter of the ripe sperm sacs of this species is about "32 mm.) Thus
these are not small polyps which would later grow into larger ones with longer tentacles
bearing more numerous and longer pinnules characteristic of some other species. They
are mature and have acquired all their adult characters.
These specimens therefore belong to a new species to which I propose to give
the name Xenia Novae Britanniae as the first specimens were obtained from Talili
Bay, New Britain (in 1895).
Diagnosis of the Species XENIA NOVAE BRITANNIAE.
The colony is knob-shaped or dome-shaped, the polyps arise from the strongly
convex upper face of the stem. The stem of the colony is usually unbranched. (See
Fig. 1.)
The measurements of the fully developed polyps are: — body 2'8 mm. — 4'5 mm. in
length (a few reach 5 mm.) and '8 mm. — 1-3 mm. in breadth, tentacles I'S mm. — 3'5 mm.
long and 7 mm. — 1"0 mm. broad. Each tentacle bears three rows of short rounded
pinnules on each side of the middle line. There are 8 — 12 pinnules in each row. The
first three or four transverse rows near the base of the tentacle are small wart-like
protuberances not more than '2 mm. long but the more distal ones consist of slightly
longer and more typical pinnules 2 mm. — 35 mm, long and 15 mm. — 2 nun. in diameter.
The pinmdes are longest near the middle of the tentacle and decrease slightly in size
towards its tip. (See Fig. 2.) There is usually in the middle line of the inner face
of each tentacle a narrow area free from pinnules which extends from the base rather
522 REPORT OX THE XEXIIDAE COLLECTED BY DR WILLEY.
more than half-way towards the tip of the tentacle. In the distal portion of the tentacle
the bases of the pinnules of the two sides are close together in the middle line. (See
Fig. 3.)
Spicules occur in considerable numbers in the body of the polyp, especially near its
distal end around the bases of the tentacles. (See Fig. 6.) They are more numerous
in the tentacles and they are so abundant on the outer face of the pinnules that they
are practically in contact with each other. (See Fig. 5.) The spicules are discs generally
oval in shape. The fully formed ones are •2mm. — -22 mm. m length, -OlS mm. — '018 mm.
in breadth, and •004 mm. in thickness. They are white or slightly bluish-white by
reflected light but reddish-brown by transmitted light.
The stem of the colony is a pale yellowish-green colour (in spirit) but the pol}7)s
are whitish with a pale bluish " bloom," due to the very numerous spicules which
they contain.
Habitat. Talili Bay, Xew Britain. (Specimens taken from trawl, 1895.)
Lifu, Loyalty Islands. (November, 1896.)
HABITS, DISTRIBUTION AND CLASSIFICATION.
The Xeniidae have been recorded fi-om the Red Sea and the tropical parts of the
Indian and Pacific Oceans. They are not recorded from the shores of America and
apparently they do not occur on the reefs near Jamaica, for although careful search
has been made on these reefs no specimens of Xenia have been hitherto found. They
are all littoral forms living in shore pools or in shallow water (3 — 6 fathoms), fixed
to the surface of coral reefs or hanging from the under side of hollow rocks or coral
boulders. They are often exposed at low water, and in those species in which the
polyps are long and slender the polyps fall together into a shapeless mass on the retreat
of the tide. They exhibit great variety and beauty of coloration; blue, green, brown,
and yellow are the predominant colours, though two species are reddish in colour.
Several of the species of Xenia founded by various authors have been subse-
quently proved to be invalid : Lamarck's Xenia purpurea is certainly not a true
Xenia but is probably a Spongodes : Schweigger's Xenia esperi and Sars' Xenia
indivisa from Naples do not belong to the genus Xenia. Duchassaing and Michelotti
described Xenia carybeorum and ,X. capitata from the West Indies, but tliese also are
not members of the genus Xenia, as they differ completely in their general characters
and in their .spicules. The former has been renamed Erythropodium carybeorum by
Kolliker.
The descriptions of several undoubted species of Xenia which have been given
are not sufficiently detailed to enable these species .to be again recognised and for
all practical purposes they must be set aside, at least for the present. Such are the
descriptions of Cornularia imdtipinnata (which is probably a true Xenia) by Quoy
and Gaimard, X. samoensis, Kolliker, X. ochracea, Sav. Kent, X. brunnea, Sav. Kent,
and X. pidsitans, Sav. Kent.
REPORT ON THE XENIIDAE COLLECTED BY DR WILLEY. 523
I have added a table of measurements and of the chief characters used in the
diagnosis of all the other known species of Xenia in the hope that it may be useful to
those who may work over the family in the future. Following Schenk the family
may be divided into three portions.
I. The tentacles of the polyps of the species forming this division bear more
or less elongated conical pinnules generally placed in two series of three rows in
each. Most species of Xenia belong to this division. (10 species.)
II. Each tentacle bears two types of pinnules, (1) smaller short round pinnules
or "warts" (Schenk) at the base of the tentacle, and (2) more typical elongated
pinnules on the more distal portion of the tentacle. (4 species.)
III. The pinnules upon the tentacles are all small conical or rounded warts
arranged either in irregular rows or scattered over the whole face of the tentacle.
(2 species.)
The numbers enclosed within parentheses in the table have been added by me
to the authors' descriptions, being taken either from the published figures or in the
cases of X. Garciae and X. coerulea from the original specimens kindly lent to me
by Mr G. C. Bourne.
GENERAL INTERNAL ANATOMY.
Sections of the stem of each, and the polyps of some, of the species above
described have been cut and examined. As all the species agree moderately closely
with A''. Hicksoni (Ashworth, 1899) in the main features of their anatomy this portion
of the report will be somewhat brief.
Stomodaeum and Mesenterial Filaments.
In many of the polyps the mouth is situated at the bottom of a slight depres-
sion '2 mm. — 4 mm. in depth, which has been produced by partial contraction of the
oral disc. The mouth leads into the stomodaeum, which is generally compressed
laterally, though in one species {X. viridis) it is almost circular in transverse section.
The stomodaeum of the fully developed polyps varies in length from "8 mm. — 2-2 mm. in
the various species. The length of the stomodaeum is doubtless related in some degree
to the length of the free portion of the polyp as in those species which possess short
polyps, e.g. X. crassa and X. Novae Bvitaimiae, in which the bodies of the polyps are
only about 3 mm. in length, the stomodaeum is only '8 nun. — •!• nun. lony. The length
of the stomodaeum does not however appear to entirely depend on the length of the
polyp, as in X. viridis, in which the polyps are comparatively short (the body measures
4 mm. — 6 mm.), the stomodaeum is 2-1 mm. — 2-2 mm. long, while in A', inembranacea. in
polyps the bodies of which are 10 mm. — 11 mm. long the stomoilacum is only 1"7 mm. —
1"9 mm. long.
In the stomodaeum of the polyps of Z. viridis and A'. Novae Britanniae, the duly two
species from this collection which have been cut into thin transverse sections, a ventral
groove or siphonoglyph is present, the cells of the lower third or two-fifths of which
bear flagella.
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REPORT ON THE XENIIDAE COLLECTED BY DR WILLEY. 525
In the same sections, among the ordinary columnar cells which line the greater part
of the stomodaeum, there are swollen and apparently empty cells similar in appearance
and position to those described in X. Hicksoni (1899, p. 251). They are cells which have
been swollen by some secretion to which they give rise, and, having discharged this
secretion, now appear empty. The stomodaeum of the stained polyps of X. crassa
also shows, when the walls are seen by transparency, numerous small light areas,
which probably indicate the presence of these secretory cells. The polj'ps of X. uni-
hellata and X. membranacea are not sufficiently well preserved to show this point in
their structure.
In all the species the polyps are devoid of ventral and lateral mesenterial
filaments but the dorsal mesenterial filaments are present and well developed. They
run in a sinuous course down the dorsal side of the coelenteron and in the primary
polyps extend to the base of the colony. (See Fig. 7.) They agree in structure with
those of X. Hicksoni.
Coelentera of Polyps.
The eight mesenteries are arranged as in typical Alcyonaria. The mesenteries of all
the species in the collection are very thin, much thinner than those of X. Hicksoni, and
their retractor muscles are feebly developed, forming only a very slight ridge on the
ventral face of each mesentery. The small size of these muscles probably accounts for
the non-retractile character of the polyps.
Between the two endodermic lamellae covering the mesentery there is a thin plate
of mesogloea which is slightly thickened near the inner or free edge of the mesentery.
In this portion cells are usually present similar to those found in a corresponding
position in the mesenteries of X. Hicksoni. These are endoderm cells which have
migrated into the mesogloea and give rise to the genital products. They have reticulate
protoplasm and large nuclei. (See Fig. 14.)
Mesogloea, its Canals and Cells.
As in X. Hicksoni, there is around each coelenteron in the stem (at any rate in
its upper part) a denser, more deeply staining cylinder of mesogloea which belongs to the
coelenteron within it. (See Fig. 7.) This is almost free from cells but is surrounded
by a cylinder of ectoderm cells in which there are numerous spicules. (1899, PI. 25,
Figs. 8, 9.) Spicules have migrated from this chain of cells into other parts of the
mesogloea, in which they are more abundant than in X. Hicksoni.
The superficial canal system is very uniform throughout, consisting of a plexus of
canals situated about '1 mm. below the ectoderm of the stem. (See Fig. 9.) The outer
wall of these canals is thicker than the inner owing to the cells of the former
being more columiuir than those of the latter wall. The cavity of this system of
canals is invaded by zooxanthellae throughout.
The internal canal system varies in the development of its parts to a considerable
extent. The extreme in one direction is reached in X. viridis, in which the longitudinal
canals which run in a sinuous course in the mesogloea about midway between the
coelentera are feebly marked. As if to compensate for this the transverse canals which
w. IV. 70
526 REPORT ON THE XENIIDAE COLLECTED BY DR WILLEY.
connect together the coelentera and the longitudinal and superficial canals are exceed-
ingly numerous, as also are the cords and strands of cells. In X. crassa and X. Novae
Bntanniae the longitudinal canals are very well marked and the transverse canals are
fewer in number than in X. viridis but still are very numerous (Fig. 8). In all species
the mesogloea (except the denser ring round each coelenteron referred to above) is
traversed by numerous strands of cells which place all its parts in intimate communi-
cation. (See Fig. 8.)
In sections which pass very obliquely through the ectoderm and mesogloea
of polyps of X. viridis small stellate cells, connected with fine fibrils which run
inwards to the endoderm, may be seen. This system of fibrils and cells has been
more fully described in X. Hicksoni (1899. p. 277), in the account of which it was
suggested that these are nervous elements homologous with the similar plexus seen
in the base of the ectoderm of Alcyonium and compared by Hicksou with the
" Nervenschicht " of the Actiniae (1895, p. 371).
Ectoderm.
In the inner or basal portions of many of the ectoderm cells of the tentacles
there are muscle fibres which are longitudinal in direction. These are much more
highly developed on the oral than on the aboral face. It is owing to their contraction
that the tentacles of some of the specimens are somewhat curled inwards over the
mouth. The ectoderm of X. membranacea contains numerous nematocysts which are,
like those of X. Hicksoni and of most other Alcyonaria, exceedingly small (see p. 513).
The spicules are similar throughout, being round or oval discs of horny consistency
(Figs. 5 and 6). On examining sections or stained preparations of the ectoderm each
spicule is seen to be accompanied by the nucleus and remains of the protoplasm of
the cell in which it has been formed.
Endoderm.
In none of the specimens does the protoplasm of the endoderm cells present the
well-marked reticulate structure which was noticed in X. Hicksoni. In A', viridis,
in which the endoderm is moderately well preserved, the cells lining the coelenteron
and particularly those covering the mesenteries are long, narrow, columnar cells which
have a finely granular protoplasm with few or no vacuoles.
Zooxanthellae are present in the endoderm of the body of the polyp but in
comparatively small numbers ; they are more numerous in the endoderm of the
tentacles and pinnules, especially in the latter, where they are sometimes so abund-
ant that they almost fill up the cavities of the pinnules.
Gonads.
Gonads are present in all the species on the edges or sides of the ventral and
lateral mesenteries, and are developed from the cells which are present in the
mesogloea of these mesenteries near their free edges (see p. 525 and Fig. 1-t). The
gonads are most abundant in the portions of the coelentera contained in the upper
part of each stem.
REPORT ON THE XENIIDAE COLLECTED BY DH WILLEY. 527
Seven colonies were examined for sexual products, five are male, one female, and
one (A", viridis) hermaphrodite, containing 3'oung ova and more advanced spermatozoa
(for more detailed description see p. 517 and Fig. 14).
The spermatozoa arise and develop similarly to those of X. Hicksoni. The ripe
spermatozoa of X. Novae Britanniae are considerably smaller than those of X. Hicksoni.
In X. uinhellata large and probably ripe ova are present. The largest ova are some-
what oval in shape and 'o mm. x "6 mm. in diameter. These ova are described in
detail on p. 515. They are rather larger than those of Alcyonium digitatum but in
other respects resemble them closely.
Addendum.
Since the completion of this Report a memoir by Walther May (Beitrdge zur
Systematik und Chorologie der Alcyonaceen. Jenaische Zeitschrift. Band xxxiii. Heft 1,
Jena, 1899) has come into my hands. This memoir contains a description of six new
species of Xenia and records the taking of four other previously known species.
May (p. 77) .suggests that probably A^ viridis, Scheuk, and A', elunc/ata, Dana,
are identical and that the descriptions of X. plicata, Schenk, and X. florida, Dana,
do not mention any characters by which the two species may be distinguished. With
regard to the former statement it may be pointed out that there are several essential
points of difference between X. viridis and X. elongata. The great length of the polyps
of the latter, shown in Dana's figure (1848, Plate 57, Fig. 5) to be 18 — 20 mm. or
even .30mm. long, and their "slender and elongate" pinnules (Dana, 1848, p. 607)
distinguish this species from X. viridis, in which the polyp bodies are ouly 4 — 7 mm.
long and the pinnules are small and thick or small round warts (Schenk, 1896, p. 62).
Dana's description of X. florida is not as detailed as we could wi.sh and there certainly
is difficulty in distinguishing this species from X. j)licata. The bodies of the polyps
of the former are of greater length, being 8 — 1:^ mm. long according to Dana's figure
(1848, Plate 57, Fig. 4) while those of X. plicata are only 4 — 5 mm. long and the
tentacles of the former are broader than those of the latter (see table, p. 524).
May also records A^ lombellata, Savigny, from Mozambique, Tumbatu and Baui
Island, X. elisabethae {= H eteroxenia Elizahethae), Kolliker, and X. membranacea, Schenk,
from Zanzibar, and X. blumi. Schenk, from Tanga and Suez.
Six new species are described by May but there are several important points with
regard to which further details would have been useful to systematists, e.g. the number
of pinnules in the rows present on the tentacles or the number of pinnules vi-sible
on one side of the outer a.spect of the tentacle of an adult polyp. The size of the
pinnules is also a useful factor. Both these characters are omitted from May's diagnoses
and descriptions. The new species are — X. tumbatuana, X. quinqueserta and X. medusoides
from Tumbatu, X. bauiana from Zanzibar and Baui Island, X. sansibariana from Zanzibar
and X. rigida from Mozambique.
70—2
528 REPORT ON THE XENIIDAE COLLECTED BY DR ^VILLEY.
LITERATURE.
1848. Dana. United States Exploring Expedition, 1838—1842. Zoophytes. 1848.
1877. Kluszinger, C. B. "Die Korallthiere des Rothen Meeres." Erster Theil. "Die
Alcyonarien und Malacodermen." Berlin, 1877.
1895. Bourne, G. C. On the Structure and Affinities of Ueliopora coerulea, Pallas, with some
Observations on the Structure of Xenia and Heteroxenia. Phil. Trans. 1895.
1895. HiCKSON, S. J. The Anatomy of Alcyoniuin digitatum. Quart. Journ. Micr. Science.
Vol. 37, Part 4, 1895.
1896. ScHENK, A. " Cla\-ulariiden, Xeniiden, und Alcyoniiden von Ternate." Frankfurt a. M.,
1896.
1899. AsHWORTH, J. H. The Structure of Xenia Hicksoni nov. sp. with some Observations on
Ueterooxnia Elizabethae, Kolliker. Quart. Journ. Micr. Sci., Vol. 42, Part 3, 1899.
EXPLANATION OF PLATES LII. AND LIIL
List or Reference Letters.
D. M. F. Dorsal mesenterial filament. Ed. Ectoderm. Ect. Ch. Chain or cylinder of
ectoderm cells surrounding the cylinder of denser mesogloea. Ect. Str. Strands of ectoderm
cells. End. Endoderm. End. Can. Endodermic canals. G. V. Germinal vesicle of ovum.
Gen. C. Genital cells in various stages of development. Lovxj. Can. Longitudinal endodermic
canal. My. Mesogloea. Mg. D. Denser cylinder of mesogloea around each coelenteron in the stem.
M. P. Muscle processes of endoderm cells. N. Nucleus. Ov. Ovum. Sp. Spicule. St. Stomodaeum.
Sup. Can. Superficial canal. S. S. Sperm sac. F. M. Ventral mesentery.
PLATE LII.
Fig. 1. Xenia Xovae Britanniae. View of a colony from Lifu (A in Table, p. 519).
The general shape of the colony, the soft, fleshy base and the strongly convex area from
which the polyps arise may be seen. The polyps are smaller and closer together near the
base of the colony, x 3.
Fig. 2. X. Novae Britanniae. One of the largest polyps of a colony from Lifu (DI in
Table, p. 519). On their outer faces the tentacles show clearly a row of pinnules on each
side, the basal three or four pinnules are small, rounded and wartlike, while the more
distal ones are longer and more typical pinnules. From the inner aspect three rows of
pinnules are visible on each side of each tentacle, x 15.
Fig. 3. X. Xovae Britanniae. A tentacle of the polyp shown in Fig. 2, seen from the
inner or oral side. Near the base of the tentacle there is a narrow area free from pinnules
separating the pinnules of the two sides. The more distal pinnules are situated close together
and many of them are flattened on one or more of their faces by mutual pressure. The
tentacle was slightly flattened in preparation and hence appears a little broader than it would
be in life, x 20.
"Wtlley. Zoolocicali Results.
Plate LII.
<S^» ;•:.•; ,-;,-', .•'A j; ty^y: ;, ;_
J.RAsWorth del.
6. t 190
West.Nevnnan lidi
ASHAATORTH. XENIIDAE .
REPORT ON THE XENIIDAE COLLECTED BY DR WILLEY. 529
Fig. i. Xenia Garciae, Bourne. One of the largest polyps of the colony, lent to me by
Mr G. C. Bourne. Note that the pinnules seen on viewing the tentacles from the outer
aspect arise close to the middle line, thus differing from those of the tentacles of X. Novae
Britanniae (compare Figs. 2 and 4). The specimen has been somewhat flattened by contact
with the sides of the bottle, x 15.
Fig. 5. Xenia Novae Britanniae. Outer surface of a pinnule of a tentacle of a polyp,
showing the very numerous disc-like spicules present in the ectoderm. The spicules are often
in contact or even overlapping, x 190.
Fig. 6. A portion of the body-wall of the same polyp 2 mm. below the bases of the
tentacles. Note that the spicules are much less numerous than in the pinnules, x 190.
Fig. 7. Xenia Novae Britanniae. A thick longitudinal section through the middle of
the colony represented in Fig. 1. Tlie strongly convex area fi-om which the free parts of
the polyps arise may be seen. The coelentera of the polyps extend a considerable distance
into the stem, those of the large polyps extending to the base of the colony. On the dorsal
side of each polyp one of the dorsal mesenterial filaments (D. M. F.) is shown and may be
traced almost to the lower end of the coelenteron ; on the ventral side of the polyp the
thin edge of the ventral mesentery {V. II.) may be seen. These have been omitted for the
sake of clearness from polyps I, II, III, VI. On the right the section passes through a
bud or young polyp (I), the free part of which is 2 mm. long. The coelenteron of this polyp
extends inlj' a comparatively short distance into the stem. The stomodaeum {St.) of two
polyps (IV, V) is shown. The coelentera of the older polyps (IV, V, VI, VII) are
crowded with sperm sacs (.S'. 8.) in the portions situated in tlie upper part of the stem,
but they have been omitted from polyps V, VI. The superficial {Sup. Can.) and longitudinal
{Long. Can.) canal systems ; their relation to each other and to the coelentera of the polyps,
the plexus of canals at the base of the colony and the ectoderm (represented when seen in
section by a thick black line) are shown. The outer walls of the superficial canals are
thicker than the inner walls. Around each coelenteron there is a denser cylinder of mesogloea
{Mg. D.), this is shown only in polyps VI, VII, being represented by the darker tone round
these coelentera. Semi-diagrammatic x 6.
PLATE LIII.
Fig. 8. Xenia Novae Britanniae. A freehand section about '15 mm. thick passing through
the column of mesogloea between two adjacent coelentera. The thick dark bands (h'nd.) riglit
and left represent the endoderm of the coelentera. The denser cylinder of mesogloea {Mg. D.)
almost devoid of cells which encloses each coelenteron, with its surrounding ectoderm cells
{Ect. Ch.): the numerous strands of ectoderm cells (Ect. Str.) passing through llie othei' portions
of the mesogloea : the longitudinal canal {Long. Can.) with its branches {End. Can.) opening into
the coelentera : the superficial canal system {Sup. Can.) seen in section with its thicker outer
walls, and its relation to the longitudinal canal system {Long. Can.) may be seen, x 70.
Fig. 9. Xenia Nocae Britanniae. A portion of the superficial canal .system whicli has
been stripped oS' the stem along with the ectoderm. The canals are nearly all in one j)lane
forming a close network about '1 mm. beneath the ectoderm, x 100.
Fig. 10. Xenia umhellata. A iiud from the edge of the sunnnit of one of the stems.
The total length of the young polyp is 3'3 mm., the tentacles are 1'5 mm. long and show,
from the outer aspect, five or six pinnules on each side of the axis of the tentacle. Viewed
530 REPORT ON THE XENIIDAE COLLECTED BY DR WILLEY.
from the inner side (see the tentacle on the left) two rows of pinnules are visible on each
side of the middle line, a third row will be formed later. This is typical of the buds of
the Xeniidae. x 15.
Fig. 11. X. umbellata. A bud from the middle portion of the summit of a stem. The
body of the young polyp has grown much larger in proportion to the tentacles than is
usual. The total length of the polyp is 6 mm., its slender linger-like tentacles are 1-.5 mm.
long and bear only one or two pinnules on each side. The tentacles of this and of the
specimen shown in Fig. 10 are equal in length. See also p. 514. x 15.
Fig. 12. A', umhellata. A bud 5 mm. long from the middle portion of the summit of
a stem. This is similar to, but probably younger than the one shown in Fig. 11. The
body is very stout, the tentacles are somewhat unequal and measure I'O mm. to 1-3 mm. in
length. Some of them (on the left) are trilobed at tlieir free ends, i.e. there is an indication
of the formation of the first two pinnules. See also p. 514. x 15.
Fig. 13. X. umbellata. Thin section (5/x) of an ovum probably almost mature. The
peripheral zone of protoi)lasm is finely granular and devoid of yolk spherules, the central
mass of protoplasm contains many cavities, which in life probably contained the }'olk spherules.
For further description see p. 515. x 200.
Fig. 14. Xenia viridis. Portion of a thin section of a mesentery which bears male
and female products. On the right the mesentery is cut almost longitudinally, showing in
the mesogloea (Mg.) the cells {Gen. C.) with well-marked nuclei and reticulate protoplasm
which give rise to genital products. In the centre of the figure are several very young
sperm sacs, each of which contains a few cells produced by division of one of the primitive
genital cells. Below and on the right is a large sperm sac containing many hundreds of
sexual cells which after a very few more divisions would give rise to spermatozoa. To the
left of the sac is one of its genital cells more highly magnified (x 2000) to show the large
nucleus (X.). On the left of the figure are four young ova (Of.) each in its follicle of
endoderm (End.). Each has a well-marked germinal vesicle ((?. V.) and germinal spot. The
small cavities in the somewhat granular pi'otoplasm probably contain yolk spherules in
life. X 320.
Erhfi
WniiEY. Zoological Resui.ts.
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ASHWORTH. XEHIIDAE.
Wf^nt NewTran Iith
/
ZOOLOGICAL EESULTS
BASED ON MATERIAL COLLECTED IN
NEW BEITAIN, NEW GUINEA, LOYALTY ISLANDS
AND ELSEWHERE.
PART V.
ILonDon: C. J. CLAY and SONS,
CAMBRIDGE UNIVERSITY PRESS WAREHOUSE,
AVE MARIA LANE,
AUD
H. K. LEWIS,
136, GOWEE STEEET. W.C.
eiasgoto: 50, WELLINGTON STEEET.
Jtfijjfig: F. A. BBOCKHAUS.
f.rto gorfe: THE MACMILLAN COMPANY.
ISDmbas: E. SEYMOUR HALE.
ZOOLOGICAL RESULTS
BASED ON MATERIAL FROM
NEW BRITAIN, NEW GUINEA, LOYALTY
ISLANDS AND ELSEWHERE,
COLLECTED
DURING THE YEAES 1895, 1896 AND 1897,
BY
ARTHUE WILLEY, D.Sc. Lond., Hon. M.A. Cantab.
LECTCRER ON BIOLOGY IN GUY's HOSPITAL, LONDON.
PART V.
{DECEMBER, 1900.
CAMBRIDGE: t\^^
AT THE UNIVERSITY PRESS. 2?^
1900
CAMBRID(jE :
PRINTED BY J. AND C. F. CLAY,
AT THE UNIVERSITY PRESS.
CONTENTS OF PAET V.
28. A Description of the Entozoa collected by Dr Willey during
his sojourn in the Western Pacific . . . . . 531
By ARTHUR E. SHIPLEY, M.A.
With Plates LIV. LV. LVI.
29. On some South Pacific Nemertines collected by Dr Willey . 569
By R. C. PUKNETT, B.A.
With Platas LVIL— LXI.
30. On the Young of the Robber Crab 585
By L. A. BORRADAILE, M.A.
With figures in the text.
31. Anatomy of Neohelia porcellana (Moseley) . . . . 591
By EDITH M. PRATT, M.Sc.
With Plates LXII. and LXIII.
32. On a new Blind Snake from Lifu, Loyalty Islands . . 603
By G. A. BOULENGER, F.R.S.
With figures in the text.
33. On Crustacea brought by Dr Willey from the South Seas .
605
By the Rev. T. R. R. STEBBIXG, F.R.S.
With Plate.s LXIV.— LXXIV.
A DESCRIPTION OF THE ENTOZOA COLLECTED BY DR WILLEY
DURING HIS SOJOURN IN THE WESTERN PACIFIC
By ARTHUR E. SHIPLEY, M.A.
Fellow and Tutor of Christ's College, Cambridge,
and University Lecturer in the Advanced Morphology of the Invertehrata.
With Plates LIV. LV. LVI.
TABLE OF CONTENTS.
Introduction
Trematoda.
Monostomum trigonocephahmi, Rudolphi .
Distommn ventricosuin, Pallas .
Cestoda.
BothrioccphaliLs plicatus, Kudolphi .
CalliobothHum. aetiobatis, n. sp.
Adelobothrium aetiobatidis, n. gen. et sp.
Palaid 7!arani, n. gen. et sp. .
Phyllobothrium dipaadomorphi, n. sp.
Coelodela kuvarki, n. gen. et sp.
Prosthecocotylc diomedeae, Fvibrmann, n. sj).
Nematoda.
Pkysaloptera obtudssima, Molin
Phymloptera return, Rudolphi .
Physaloptera varani, Parona
Echinocephalus striahis, Monticclli .
Sderostomum appendiculaium, Molin
Gixathoitonia shipleyi, Stossich
W. V.
PAGE
532
532
540
540
541
545
548
550
552
557
559
559
560
560
560
560
71
532 A DESCRIPTION OF THE ENTOZOA COLLECTED BY DR WILLEY
PAGE
Nematomorpha.
Oordius willeyi, Camerano "^^
LiNGUATULIDAE.
Porocephaliis tortus, Shipley 5"^
INTRODUCTION.
The collection of parasites brought back by Dr Willey from the far East was a
large and comprehensive one. If we include a small tick from the skin of the
"Malagea" snake Dipsadomorphus irregularis Merrem, which I have unfortunately failed
to get identified, the collection contains examples of all the groups of Metazoa which
infest land animals, except the Acanthocephala, and practically of all those, with the
exception of the Crustacea, which are found in or on marine creatures.
The collection may be briefly summarized as follows : Trematodes, two species ;
Cestodes, seven species, including three new genera and six new species ; Nematodes,
six species, one of them new ; Nematomorpha or Gordian worms, one species, new ;
Pentastomids or Linguatulids, one species, new.
I have as far as possible given the British Museum Catalogue names to the hosts,
and have throughout, in referring to journals and periodicals, used the abbreviations
which are suggested by Mr D. Sharp in the Zoological Record.
I. TREMATODA.
I. MONOSTOMUM TRIGONOCEPHALUM Rud.
The following brief account of the literature precedes a detailed description of the
anatomy of this form.
O'B. Bellingham. Ann. Nat. Hist. (i. Ser.), Vol. xiii. 1844, p. 337.
The author has not found this parasite nor does he give any authority for in-
cluding it in his " Catalogue of Irish Entozoa," but he mentions that its host has
been occasionally thrown upon the English coast, so that the parasite has some " claim
to be considered British." Let us hope however that not every turtle is infested with
these Trematodes.
C. M. DiESlNG. Systema Helminthum. Vienna, 18.50, Vol. I. p. 325.
References will be found here to the older literature and to the synonyms.
P.-J. VAN Bkneden. Bull. Ac. Belgique (ii. Ser.), Vol. VL 18.59, p. 81.
This author appears to doubt if the Trematodes seen by Van Hasselt and Kuhl
and described by them under the names Monostoma rubrum and Monostoma album are
DURING HIS SOJOURN IN THE WESTERN PACIFIC. 533
the same as Rudolphi's species. Diesing however includes these amongst the synonyms
of Monostomum trigonocephalum.
E. Walter. Zeitschr. wiss. Zool. Vol. 56, 1893, p. 189.
This paper includes a comparatively complete account of M. proteus Brandes and
M. reticulare van Ben., but the account of M. trigonocephalum is more meagre, so that
I have thought it worth while to print the following notes on the anatomy of this
form, but for many details I must refer the reader to Walter's paper.
G. Brandes. Centrbl. Bakter. Vol. xii. 1892, p. .505.
A. Looss. Zool. Jahrb. Syst. Vol. xii. 1899, p. 523.
External Features, Ectoderm and Parenchyma.
Since this article has been in type,' Professor A. Looss has published a most
valuable essay on the Trematodes of Egypt. He gives a short account of M. tiigono-
cephalum under the name Pronocephalus trigonocephalus {= M. trigonocephalum Rud.
partim) and a figure. With the exception of a few details his description agrees
with the following.
Dr Willey brought back with him some portions of the intestine of a Ghelone imhri-
cata L. attached to which and half hidden in the mucous secretions were a small number
—some ten or twelve — Trematodes belonging to Rudolphi's species Monostomum trigono-
cephalum. The shape of these Trematodes varied markedly (Fig. la, lb, 1 c, Id,
Plate LIV.), some had an almost U shaped venti-al flexure, others were straight, but
all possessed the marked ventral hollow which caused van Beneden to describe the
species " excave comme un canot." The longest specimen would have measured some-
thing over 5 ram. had it been straightened.
The body is covered by a cuticle of uniform thickness, this is apparently secreted
by extensions of certain cells which I take to be ectoderm and whose body with the
deeply stained nuclei are withdrawn some distance from the surface. Amongst these
cells run longitudinal and oblique fibres (Figs. 2 and 3, Plate LIV.) but I did not find
any absolutely transverse fibres : perhaps they are replaced by the diagonal fibres, which
are after all only slightly out of the transverse plane. Dorso-ventral fibres cross the
body in a vertical direction and are inserted above and below into the cuticle. They
often indent various organs, e.g. the testes, so as to destroy the evenness of their
surface. The solid parenchyma of the body consists for the most part of large cells
which tend to become very vacuolated (Fig. 3, Plate LIV.). These cells have minute
nuclei and the protoplasm stains very slightly. Every now and then a more compact
cell, better stained and with a larger nucleus, is met with embedded in the large
celled parenchyma. Such cells are especially common near the head where the looser
vacuolated cells are almost absent (Fig. 7).
71—2
534
A DESCRIPTION OF THE ENTOZOA COLLECTED BY DR WILLEY
The Reproductive System.
One or two points in the
Yftds
Fig. a. Diagram of the reproductive
organs of Jilonostoiiutm trigono-
cephalum Rud.
Egg car. egg cavity, Gen. sin. geni-
tal sinus, Glds. glands at base of penis,
Ov. ovary, Sc. sucker, Sh. ghi. shell-
gland complex, Tl: right testis, Ut.
uterus, Vag. vagina, Vas. def. vas
deferens, Ves. sem. vesicula seminalis,
r. glds. yolk glands, .-1. rod-shaped
gland, B. dome - shaped organ, C.
glands opening into vagina.
Reproductive System have hitherto been overlooked and
as the system is very complex I think a detailed description
will not be out of place.
Male. The testes are two in number and lie at the
same level near the posterior end of the body (Fig. A)', and
as Walter points out their presence causes the forks of
the alimentary canal to bend in towards one another near
theu- posterior terminations. All stages in the spermato-
genesis can be seen in the cells of the testes. These organs
have no muscular walls and seem to lie freely in the
parenchyma. Each testis is roughly speaking spherical in
shape but the dorso-ventral muscles press into it in places
and thus in some sections it seems to be lobed (Fig. .5,
Plate LI v.).
From each testis a short vas efiferens passes inwards and
forwards and verj- soon meets its fellow, and the two fuse
into a median vas deferens. This is as a rule filled with
spermatozoa. As it passes forward the vas deferens widens
and passes without an abrupt change into a capacious thin
walled vesicula seminalis which curves from side to side in
the most dorsal part of the body (Fig. A in text). The
vesicula is crowded with spermatozoa which as a mass
stain deeply, they are matted together like -ivisps of hay
and I could detect no head, nothing but a long tail.
About half-way up the body the vesicula seminalis
passes into a very remarkable organ, which as far as I
can gather has not hitherto received much notice, possi-
bly it is only present in those specimens which are sexually
mature. Like the vesicula this organ i.s situated just
below the dorsal skin and it is median and unpaired. It
consists of a hollow rod-like gland with a well-developed
lumen (Fig. 4, Plate LIV.). Internally the gland is lined
by large cells which stain very deeply. The free ends
of these cells project like little balloons into the cavity
of the organ, and these swellings although ci'owded with
granules do not stain so deeply as the body of the cells.
1 Looss states that "Die Hoden liegen bei meinen Exemplaren ausuahmlos asymmetrisch schrag hinter
einander, nur bei sehx stark contrahirten Thieren kommt gelegentlich der An sc he in einer symmetrischen
Lagerung zu Stande."
I had no opportunity of observing the animals alive and my specimens had undoubtedly contracted, and
in them the right and left testes are situated at practically the same level.
DURING HIS SOJOURN IN THE WESTERN PACIFIC. 535
From the way these ends project a section passing in the plane of the inner face of
■ the wall of the organ cuts them, so that they appear to lie freely in the lumen of
the gland. This may account for many of the vesicles which help to fill the lumen
but I think not for all. Some of the vesicles have I think left the cells which
produced them and lie in the lumen of the gland. Outside the deeply stained basal
ends of these cells is another layer of large cells possibly destined to grow up into
the glandular inner cells, and still further outside is a well-developed stout layer of
longitudinal muscles. The muscle-fibres building up this layer are arranged in bundles,
and each fibre is tubular with a lightly stained contents and closely resembles the
fibres in the Leeches. They seem to be more specialized than any other muscle-fibres
in the animal.
The vesicula seminalis opens by a small opening into the posterior extremity of
this organ, but I have not seen, in any of the six specimens I cut, any spermatozoa
inside the organ and indeed the anterior end of the vesicula is free from them.
In front this organ opens into a dome-shaped chamber. Between the two is stretched
a diaphragm and in the middle of this is a beautifully neat little funnel-shaped valve
with circular muscles in its walls (B, Fig. 4, Plate LIV. and Fig. A in text).
I find it difficult to suggest any explanation of this organ, it may however
possibly be that its secretion is used to build up the spermatozoa into spermatophores.
If this be the case they must be extremely small as the lumen of the next part
of the male duct is very minute. Sedgwick' has suggested that in Trematodes, "It
may be that in some cases the penis is used, like that of Turbellaria, for hypodermic
injection of spermatozoa," and the organ of such hypodermic injection is replaced in
higher forms, i.e. Leeches and possibly Peripatus by spermatophores, but I am unaware
that these structures are known amongst Trematodes and even if they are the view
that they are formed by this enigmatical organ is highly conjectural.
One feature is of interest and that is the mode by which the secretion leaves
the cell which elaborates it. Similar modes of secretion in which the secreta are
collected in a vacuole at the free end of an epithelial cell are well known in the
mammary gland, in the digestive glands of Crustacea, e.g. Astacus, and has been
described by myself in the nephridia — brown tubes — of the Gephyrea^ It is probably
more widely spread in the animal kingdom than is usually recognized.
The dome-shaped chamber was as a rule empty but in one specimen it contained
a mass of minute granules in appearance resembling those of the vesicles but no
longer retained in vesicles. Scattered amongst these were a few spherical corpuscles
of unknown provenance, which stained very deeply. At its apex the dome-shaped
chamber opens by a small pore into the minute duct of the penis. This duct is at
first almost capillary and is contained in a muscular sheath with fibres running both
longitudinally and circularly. The tube has a cuticular lining and as it twists in the
substance of the penis and the penis is itself much coiled the lumen of the tube
appears many times in each section. After a course of some length this minute tube
widens, its walls cease to show the definite cuticular lining and they become relatively
' .-1 Htudent's Text-Book of ZooUnjij. London, 1898.
= Quart. J. Micr. Sci., Vol. xxxi. 1890, p. 1, and Vol. xxxii. 1891, p. 111.
536 A DESCRIPTION OF THE ENTOZOA COLLECTED BY DR WILLEY
thinner though they retain their muscular character. The lining of this part of the
penis usually shows a frayed margin something like a ciliated surface but I doubt"
if the appearance is due to cilia. The lumen in many sections is semi-lunar shaped,
the presence of a typhlosole reducing it to a curved slit (Fig. 5, Plate LIV.). All
along the outermost portion of the penis the tube receives the secretion of numerous
glandular cells more or less aggregated into a gland. The penis curves to the left
of the "egg-cavity" and opens into a very shallow small genital sinus a little to the
left of the middle line not very far behind the posterior limit of the pharynx (Fig. A
in text).
Female. The female organs are equally complicated. The ovary is single, dorsal,
posterior and slightly to the left of the middle line. It consists of compact cells, the
ova ; the less mature show a very well marked chromatin tangle in the middle of their
nuclei. The ripe ova have that curious cubical look with rounded corners which is
characteristic of many Trematode eggs. The oviduct is short and opens almost directly
into the shell-gland complex.
The yolk glands are densely staining compact glands lying ventral to the testes,
stretching in front of that organ for about a third of the way or perhaps one half
the way along the body. They consist of cells apparently crowded with yolk granules.
Their ducts, one on each side, converge and enter the shell-gland complex (Fig. A in
text).
The shell-gland is median, it consists of numerous glandular cells surrounding and
for the great part forming the walls of the structure which I have referred to as
the shell-gland complex. I have not succeeded in tracing out all the details of this
minute organ. I have traced the oviducts, and the ducts of the yolk glands into a
cellular complex with a canal. The canal is I believe continuous with the very short
oviduct and its cellular walls the shell-glands. From this I have traced the beginning
of the uterus with young eggs in it, but I can offer no explanation as to how the small
ovum with its large nucleus is packed up with the yolk granules and each surrounded
by its cuticular egg-shell. It must however all take place in this minute organ.
The lower end of the oviduct soon widens until we find a tube with thin walls
and a uniform lumen which passes from side to side of the body forming some thirty
loops (Fig. 5, Plate LIV.). These are very much simplified in the diagi-am (Fig. A
in text).
Towards the anterior end of the body this oviduct opens into an irregular space
which I have called the " egg-cavity." Its walls stain deeply and when I first saw
it in section I took it for a part of the ventral hollow which projects so far into
the under.side of the body. It however has no communication with the exterior except
through the vagina. The latter leaves its anterior end and is at first a fairly wide
tube. During its course two similar and highly peculiar organs open into it, one nearer
the inner and the other nearer the outer opening.
Each of these organs has a somewhat Hattened spherical shape, one surface pro-
jecting freely into the lumen of the vagina. The substance of the organ is homogeneous,
no cells can be detected in it and the whole stains lightly but uniformly. Scattered
DURING HIS SOJOURN IN THE WESTERN PACIFIC. 537
through this uniform mass are a number of deeply stained structures shaped like pins
with rather big heads (0, Fig. A in text). The pin is embedded in the mass, the
head deepest and the point coming to the free surface on which it apparently opens.
Sections of these structures are shown in Fig. 7, Plate LIV., the largest was unfortu-
nately fractured in cutting. I can offer no suggestion as to the nature or meaning of
these organs.
An occasional egg is seen in the vagina, the lumen of which is bounded by a
frayed deeply staining wall recalling that of the penis, and as in that case it may
be that this appearance is due to cilia. The vagina is surrounded by a number of
unicellular glands massed together into follicles, which closely resemble in structure and
appearance the glands surrounding the penis in the male.
Professor Braun in the article on Trematodes in Bronn's " Thierreich " says that
a Laurer's canal is usually absent in the family Monostomidae and is absent in the
genus Monostomum. I have not succeeded in seeing one in my sections but Walker
has found the canal in all three species he worked at, and in some of my sections the
inner end of the uterus and the shell-gland complex was full of eggs and spermatozoa
mixed ; it seems more in accordance with what is known of the group that these
should have entered by a Laurer's canal than have made their way up the complicated
female passages. No spermatozoa were seen in any other part of the female apparatus
in my specimens.
The eggs undergo considerable changes between the time they leave the shell-
gland complex and when they leave the body. The youngest egg in the uterus is an
oval body containing four or live yolk granules quite discrete and distinct, and
arranged in a row (Fig. 6, Plate LIV.). These all stained deeply, and one of them
may have been the nucleus which otherwise was indistinguishable. The protoplasm
stained hardly at all in this .stage. The egg is presumably fertilized before the
shell was deposited. This shell is at first a thin cuticular covering apparently
permeable by staining fluids; as the eggs pass down the uterus the shell thickens,
and it soon appeai-s with a very distinct double contour. The staining fluid no longer
penetrated the thick shell readily and only in those sections stained on the slide did
the protoplasm take up any pigment. When this was done however the egg took up
the colouring matter uniformly and the yolk granules no longer appeared as separate
deeply stained bodies (Fig. 7, Plate LIV.).
At first the eggs are oval, but about half-way along the uterus a stalk begins
to appear. These seem to increase in length — I do not know how — until they are
some four or five times the length of the egg. They are not quite stiff but can be
bent and sometimes are curved by being pressed against the edge of the uterus,
usually however they are quite straight and lie together in parallel bundles recalling
the lances in Velasquez's "Surrender of Breda." I never saw an egg with two filaments
arising from opposite poles, as Walter figures them, and I never saw the filaments
pointed, they always ended squarely, but he mentions that the form of these stalks
is very variable and my observations serve to confirm the truth of his last sentence
"Demnach niussen wir annehmen, dass die Ausbildung der Eifilainente bei dieser Species
variirt."
538
A DESCRIPTIOX OF THE ENTOZOA COLLECTED BY DR WILLEY
The Alimextary Caxal.
Ceph.gaAg.
The mouth lies in the middle of the sucker which is terminal, and opens on
the ventral face (Fig. 3, Plate LIV.). The buccal ca\'ity is triradiate, one angle pointing
backwards and leading into the narrow oesophagus. Amongst the radiating and circular
muscle-fibres in the walls of the suckers were scattered some spherical cells which
may have been glandular, but in my opiaion are more probably connective tissue.
The fine lumen of the oesophagus is lined by a cuticle surrounded by a circular and
then a longitudinal muscle-sheath.
After a very short course the oesophagus divides into two branches, each of
which runs towards the edge of the animal and then
turns backward, forming one limb of the forked gut
(Fig. B in text). The numerous shallow diverticula of
these branches and the histological details are fully de-
scribed by Walter. Each limb of the gut is somewhat
crinkled in its outline. An important point which is
shown in the diagram is that the lateral diverticula are
all on the outer side of each limb, none pass inwards
between the two limbs. They are very regularly ar-
ranged and lie in the same plane. The cells lining
each limb and the diverticula are low epithelial cells
with small nuclei and numerous vacuoles. Owing to
the irregularity of the outline of the walls of the
alimentary canal its lumen, which in my specimens
was always empty, is roughly star-shaped.
Nervous System.
Tucked into the angle between the globular pharynx
and the slender oesophagus are two half-moon shaped
nervous masses (Fig. 3, Plate LIV. and Fig. B in text).
They consist of nerve fibrils and ganglion-cells, the
latter as a rule nearer the periphery. Slender cords
connect the mass dorsally above the oesophagus. Yen-
trally and anteriorly a short nerve is given off on
each side which runs along the lower surface of the
pharynx and soon diwippeai-s. Yentrally and posteriorly
a stouter nerve passes backward on each side and nins
along the ventral surface of the animal just within the
epithelium, giving off transverse branches both exter-
nallv and internallv.
' si. Can
Fig. B. Diagram of the alimentary
canal, nervous sj'stem and water-
vascular system of Monostomum
trigonocephalum Rud.
Al. Can. branches of alimentary
canal, Ceph. gang, cephalic ganglion,
Ex. Can. lateral excretory canal, water-
vascular vessel, N. nerve cord, Sc.
sucker, v. vesicle opening at pore of
Ex. Can.
DURING HIS SOJOURN IN THE WESTERN PACIFIC. 539
Walter, who gives a much more complete account of the nervous system of
M. proteus, an animal better adapted for the investigation of this tissue than is
M. triyonocephulum, describes nerves going off from the side of the ganglia in addition
to the two mentioned above. I was only able to find such a nerve on the left side,
it soon lost itself in the outer end of the generative organs.
There is no connective tissue sheath to the nervous system, which lies simply
embedded in the parenchyma.
The Excretory System.
The excretory pore is situated at the posterior end of the body, behind the
hindermost parts of any of the other systems of organs, yet it is not quite terminal
but rather opens on the dorsal side of the body (Fig. 8, Plate LIV.).
The pore is median and minute, it leads into a very tine duct lined with a stout
cuticle, the mother-cells of which are small with large nuclei and these stand out
in marked contrast to the pale parenchyma which surrounds them. Passing forward
this median duct soon reaches the level of the terminal branches of the alimentary
caeca and here it enlarges into a capacious vesicle. The vesicle is posteriorly surrounded
by cells similar to those which encircle the duct, anteriorly however these cells are
absent and all tbat is to be seen in section is a clear space surrounded by a sharply
defined cuticle which externally abuts on the parenchyma of the body.
Walter describes in M. proteus and in M. reticulare, ceitain ciliated grooves or
diverticula which form the posterior walls of the vesicle and which converge towards
the pore. A similar funnel-shaped opening exists in M. trigonocephalum but Walter
is not so clear as to the ciliation of its ftirrows and ridges. I also have seen
appearances which may be due to cilia badly preserved but before pronouncing an
opinion I should like to see them flicker in a live animal. Such a ciliated external
termination of the excretory system is as for as I know not described in other genera
of Trematodes. It may obviously help to expel the waste products and its presence
may be correlated with the absence or very slight development of the musculature
of the vesicle which is a characteristic feature of the three species mentioned above.
About the level of the anterior third of the testes the vesicle, which by this
time has narrowed, splits into two main tubes which continue to run forward in the
same horizontal plane and at first lie parallel one to another.
Each branch runs forward, lying on the whole below the limb of the alimentary
canal of its side. Anteriorly the right and left branch unite below the oesophagus.
About half-way along the length of the chief lateral vessel — the exact position is not
invariably the same in different .specimens — a secondary vessel arises which passes still
more ventralwards and runs in the edge of the animal to the extreme posterior limit
of the body (Fig. B in text).
I could trace no smaller vessels arising from these main ones and could see
w. V. 72
540 A DESCRIPTION OF THE ENTOZOA COLLECTED BY DR WILLEY
nothing of any flame-cells. The walls of the vessels are lined with a thin cuticle
on which rests an occasional nucleus. The most interesting feature apart from the
ciliated excretory vesicle, if the cilia really exist, is the presence of numerous recesses
or globular diverticula along the whole course of the vessels (Fig. 7, Plate LIV.). In
sections these diverticula give the idea that the vessel is splitting into branches but
as one passes on to the next two or three sections it becomes evident that no
branching is taking place but that an almost spherical bulging of the wall of the
tube has occurred. The rim which lies between these globular diverticula and the
lumen of the vessels is thickened at the edge and presents the appearance of a valve
such as has frequently been described in the excretory tubules of the Cestoda
<Fig. 7, Plate LIV.).
II. DISTOMUM VENTRICOSUM Pallas var. minor.
G. R. Wagexer. Arch, Naturg. 26th Jahrgang, I. 1860, p. 166.
R. MoLiN. Denk. Ak. Wien, xix. 1861, p. 209 (D. ocreatum).
P.-J. VAN Beneden. Mem. Ac. Belgique, xxxviii. 1871, Les Poissons des Cotes de
Belgique, p. 68.
This parasite, for whose identification I am indebted to Professor M. Stossich of
Trieste, has hitherto been found in certain species of Clupeidae. Dr Willey took his
specimens from the stomach of a Sea-bream, Pinielepterus sp. in New Britain.
11. CESTODA.
I. BOTHRIOCEPHALUS PLICATUS Rud.
C. M. DiESiNG. Systema Helminthum, Vindob, 18.50 — 1851, p. 591.
This work contains a list of synonyms and references to literature anterior to the
year 1850.
G. R. Wagener. Acta Ac. German. Vol. xxiv., Supplement, 1854, p. 71, PI. VIII.
figs. 94 and 95.
T. S. CoBBOLD. J. Linn. Soc. Vol. IX. 1868, p. 200.
P. Olsson. Acta Univ. Lund. Vol. iv. 1867—8, Article VIIL, p. 11.
Dr Willey brought home with him some portions of the intestine of the sword-
fish found in the Indian and Pacific Oceans, Histiophorus sp. which were crowded
with a Tape-worm. This I take to be the species Bothriocephalus plicatus of Rudolphi
recorded from Xiphias gladius L., the better known sword-fish.
It is unfortunate that it was impossible to determine the species of the host
but as Giinther' remarks, "The distinction of the species is beset with great difficulties,
' An Introduction to the Study of Fishes. Edinburgh, 1880.
DURING HIS SOJOURN IN THE WESTERN PACIFIC. 541
owiug to the fact that few examples exist in museums, and further because the form
of the dorsal fin, the length of the ventrals, the shape and length of the sword
appear to change according to the age of the individuals."
As far as I am aware no Entozoa have as yet been recorded from this eastern
sword-fish, the genus is not mentioned in von Linstow's " Compendium der Helmintho-
logie " and I have met with no reference to this fish as a host in later literature.
The fish is in fact in the adult stage difficult to capture, it is " the largest of
Acanthopterygians and not exceeded in size by any other Teleostean " and is unusually
strong and fierce.
The most complete diagnosis of this species B. plicatus is that given by Olsson,
which I here copy.
" Caput ovato-oblongum, apice depressuiii, hothriis oblongis laterulibus. Collurn nullum.
Articuli brevissiini, sensiin latiores, murginibus posticis exstantibus, plerumque undulatim
crispatis ; aperturae genitalium (?) laterales, marginibus approximatae, vage alternae.
Long. 350"""-, lat. max 17, lat. minima 2"""-, long, ovuli Oil"""."
None of the specimens collected by Dr Willey attained the length of 350 mm. or
the breadth of 17 mm. The longest fragment was some 200 mm. and the widest segments
were by 3 or 4 mm. broad (Fig. 10, Plate LV.). They break with ease and I have
little doubt that if examined alive longer forms would have been found.
As a rule the head and anterior end of the body are embedded in the mucus
lining the intestine, but they are readily detached, the head being but very slightly
adherent to the tissues of the host. The anterior end of the head bears a flattened
cap divided into four simple lobes (Figs. 8 and 9, Plate LIV.). In the centre of this
a slight projection is sometimes seen and is figured by Wagener though it is often
absent, but I could not agi-ee to the statement " caput... apice depressum" as universally
true.
The two long slit-like suckers which extend along the head below the cap are
situated in a plane which corresponds with the line separating the two lobes of the
right side from the two of the left.
II. CALLIOBOTHRIUM AETIOBATIS, n. sp.
In his admirable paper entitled "A Cz^paknak es Rajaknak Belfergei'" the late
Dr Orley gives the following systematic review of the genera of tape-worms found in
the Elasmobranchii. I have ventured to copy the table because, although the Latiu
leaves something to be desired, it gives a clear view of the inter-relation of the
Cestode genera which infest Selachians and because the Hungarian periodical in which
it appears is somewhat difficult of access.
' Termfszetrajzi Fiixetek, Vol. ix. 1884, p. 97. Budapest.
72—5
542 A DESCRIPTION OF THE EXTOZOA COLLECTED BY DR WILLEY
CONSPECTUS DISPOSITIONIS SELACHIORUM ENTOZOORUM.
Ordo : Cestoidea Rud.
1810. Eatozoorum (45), Vol. ii. P. ii. p. 3.
SuBORDO : Paramecocotylea Diesing.
1850. Revision (22), p. 11.
Fam. (a) Tetraphyllidae van Beneden.
1850. Recherches (7), p. 112.
Caput bothriis quatuor sumrae versatilibus instructo.
I. Bothria inermia Phyllobothridae.
II. Bothria uncinulis armatis Phyllacanthina.
III. Bothria proboscidibus terebratoriis armatis, in
coUum retractilibus instructum Phyllorhynchidea.
I. SuBFAM. Phyllobothridea van Beneden.
1850. Recherches (7), p. 113.
A. Corpus dense aculeatum.
1. Genus. Cylindrophorus Diesing.
B. Corpus nudum.
(a) Bothriis quatuor pedicello contractili affixis.
2. Genus. Anthobothrium van Beneden. Bothriis indivisis.
3. Genus. Echeneibothriuvi van Ben. Bothriis transverse costatoplicatis.
4. Genus. Orygmatohothrium Diesing. Bothriis quatuor, singulo acetabulis aux-
iliaris instructo.
(6) Bothria capiti adnata (Bothria sessilia).
5. Genus. Tetrabothrium Rud. Bothriis indivisis; acet. auxil. nulla.
6. Genus. Monorygma Diesing. Bothriis indivisis; acet. aux. instructis.
7. Genus. Phyllobotfn~ium van Ben. Bothriis quatuor, marginibus laciniato-
crispatis, singulo acet. aux. instructo.
8. Genus. Trilocidaria Olsson. Bothriis quatuor, singula bothi-ia loculis ternis
in triaiigulum dispositis.
DURING HIS SOJOURN IN THE WESTERN PACIFIC. 543
II. SuBFAM. Phyllocanthina van Ben.
1850. Recherches (7), p. 127.
A. Bothriis quatuor, singulo uncinalis 2, magis curvatis, bifurcatis armato.
9. Genus. Acanthohothnum van Ben.
'B. Bothriis quatuor, singulo uncinali.s 4, simplicibus, parum curvatis armato.
10. Genus. Onchobothrium Rud.
11. Genus. Calliobothrium van Ben.
III. SuBFAM. Phyllorhynchidea van Ben.
1850. Recherches (7), p. 14-i.
A. Bothria nulla.
12. Genus. Abothros Welch.
B. Caput bothriis instructis.
13. Genus. Rhynchobothrium Rud. Caput bothriis duobus oppositis parallelis
aut apice convergentibus.
14. Genus. Tetrarhynchus Rud. Caput bothriis quatuor, binis oppositis, capiti
parallelis.
Familia(6): Diphyllidea van Ben.
1850. Recherches (7), p. 158.
Caput versatile bothriis duobus lateralibus et infra marginem anticum utrinque
rostello horizontali protractili, apice armato munitum.
15. Genus. Echinobothniim van Ben.
A point of interest in the Helminthology of Elasmobranchs is the minuteness of
the parasites. As Dr Orley records, the tape-worms which infest the largest sharks
such as Carcharias and Heptanclius never surpass 10 cms. in length. As a rule the
size of the Entozoon is inversely proportional to that of its Elasmobranch host. Both
the small .size and the comparative rarity of the parasites point to the fact that
Elasmobranchs suffer but little from the presence of Cestodes, though doubtless the
Teleosteans in whose bodies for the most part the worms pass through its cystic stage
sufifer considerably. When however they do occur in Elasmobranchs they are often met
with in great numbers, and this is especially the case with the genus Calliobothrium
which is sometimes found in enormous numbers on the spiral valve.
In that helminthological mine of information P.-J. van Beneden's " Recherches sur
la Faune Littorale dc Belgicpio, Les Vers Cestoides,'" the author establishes the genus
' Mem. Ac. lieliiiqitr, xxv. 18-50.
544 A DESCRIPTION OF THE ENTOZOA COLLECTED BY DR WILLEY
Galliohothrinm and describes the three species C. verticUlatum Rud. from Mustelus
vulgaris and also from Galeus canis and Squatina angelus. C. leuckartii from Mustelus
vulgaris and G. eschrichtii from the same host. Zschokke in his Monograph '"' Recherches
sur la Structure Anatomique et Histologique des Cestodes,"' describes C. filicolle from
the spiral valve of a Torpedo marmorata, G. crassicole from the same organ in a
Trygon pastinaca, and G. uncinatum found in the intestine of Raja batis and Haja
clavata and Trygon pastinaca. A seventh species G. coronatum is described in P. -J.
van Beneden's " Les Poissons des Cotes de Belgique "' and by Olsson in his " Entozoa,
iakttagna hos Skandinaviska hafsfiskar"^ from Eajn batis and Raja clavata, ScyUium
catulus and ScyUium stellare.
In an Aetiobatis narina?-i Euphras. taken at Lifii, Dr Willey found two species
of Cestoda, both living in the middle region of the intestine where the spiral valve
lies. The extreme upper end of the intestine was free from parasites, the lower end
contained no Cestodes but some Nematodes. The species which I have described below
as Adelobothriuin was much more plentiful than the other which proved to be a new
species of Galliobothrimn. The latter svas easily removed but the former was firmly
attached to the walls of the intestine and was i-emoved with difficulty. No hooks
could be discovered on this species and suckers could not be detected.
There can be no doubt that the less numerous Tape-worm found in the Aetio-
batis narinari belongs to the genus Galliobothrium. Zschokke* points out that " Le
genre Galliobothrium serait caracterise sufiisamment par le seolex portant quatre both-
ridies divises invariablement en trois compartiments superposes par deux hourrelets
transversaux. Le sommet du seolex porte quatre ventouses auxiliaires plus ou moins
developees. Au bord superieur de chaque bothridium on trouve : ou bieu quatre
crochets simples (C. Leuckartii et 0.^ verticillatum) ou deux crochets bifurques, (C.
coronatum, crassicolle, uncinatum, Jillicolle). En tout cas le seolex est armd de seize
pointes."
My species has the hooks bifurcated and therefore is more nearly allied, as
to this point, with Zschokke's second group. In sections it becomes apparent that
the hooks are hollow at their base like the horn of a hollow horned ruminant. I have
carefully compared the animal with figures of the above-mentioned species and find
that it differs from them in certain points which I think are of specific import-
ance.
In all the figui'es and descriptions of species of Galliobothrium that I have seen
the greatest diameter of the head is either about half-way between its anterior and
posterior faces or at its posterior end. In the former case the head is barrel-shaped,
in the latter it resembles a pear. In the species in question, however, the head is
very definitely broader at its anterior surface ; it narrows in the centre and expands
again posteriorly though not to quite the same extent as in front (Fig. 12, Plate LV.).
' Mem. Iivit. Geiiev. xvii. 1889.
- Mem. Ac. Belriique, xxxviii. 1871.
» Acta Unic. Lund. m. 1866—7.
■• Mem. Inst. Genev. xvii. 1886— 188il.
" Zschokke unites the genus Onchobothrium with Culliobntltrium.
DURING HIS SOJOURN IN THE WESTERN PACIFIC. 545
Roughly speaking the head is hour-glass shaped or perhaps it more closely resembles
a box for throwing dice.
The length of the head is a trifle under 1 mm. and its breadth averages about
half its length, the length of the whole animal is from 20 to 30 mm. The pro-
glottides are flattened anteriorly, but posteriorly they swell out and assume a some-
what bolster-like appearance (Fig. 11, Plate LV.). They are in no part separated one
from another by deep furrows and there is no projecting lip overlapping their anterior
end.
The four primary suckers which lie on the anterior face at the base of the hooks
are round (Fig. 12, Plate LV.). The third bothriuni is very much smaller than the two
which lie in front of it and is separated from the second by a very slight ridge.
The water vascular system makes a mesh-work in the head which reaches to the
extreme anterior limit. Both ventral and dorsal cord persist through the mature
proglottides but the ventral one is a good deal larger than the dorsal one.
The parenchyma of the body is loose and in places takes up very little stain
and has a clear appearance.
The reproductive organs correspond in all essentials with those described by Zschokke
in other species of the same genus. The lumen of the vagina is lined by what look
like cilia and it is unusually spacious. The cirrhus is provided with rows of large
spines, much larger than those in any other species of which I have seen figures.
The specimens at my disposal were too few in number to make sections in all
planes of the head and I have been unable to work out the details of the muscles
in this region. Certain of the longitudinal muscles however pass from the head into
the neck and these are arranged in eight symmetrical bundles, each bundle seems to
lie in a sheath and the whole i-ecalls a somewhat similar structure in Tetrarhynchus.
The following is a short diagnosis of the new species :
CalUohothrium aetiobatis, sp. nov.
This species is characterized by the shape of the head, which is like a dice-box,
narrow in the middle and broader at each end. The hooks are bifurcated and hollow
at the base. The posterior bothrium is very small and only separated from the
middle one by a very insignificant lip. The suckers on the anterior face of the head
are circular, and the muscles running from the head through the neck are eight in
number and lie in sacs. The penis is armed by unusually large spines, which are
slightly curved.
Habit.\T. Found in the intestine of Aetiobatis imrinari Euphras. in the region of
the spiral valve.
III. ADELOBOTHRIUM AETIOBATIDIS, n. gen. et sp.
Of the two tape-worms which were found in the intestine of Aetiobatis narinari
Euphras., Adelobothrium aetiobatidis is by far the commoner. I have spent a consider-
able time in trying to identify this Cestode with some already described form, but
neither in the writings of P.-J. van Beneden, Orley, Zschokke or Linton who describes
546 A DESCRIPTION OF THE ENTOZOA COLLECTED BY DR WILLEY
forty-two species and twenty-two genera of Cestodes from Fish or in Bronn's " Thierreich "
have I been able to find anything which I could regard as congeneric with the form
which I have ventured to name as above.
The length of the animal, with ripe proglottides, as shown in Fig. 13, Plate LV.,
averages about 4 to 5 cms., its breadth in the broadest part 1"5 mm. The grooves
dividing the anterior segments are deep, so that the edges project as conspicuous
flaps in the middle region of the body, the edges overlap to a very marked extent
the succeeding segments so that a transverse section usually cuts a given segment
where it is surrounded by the overlapping lips of the hinder border of the pre-
ceding proglottis. The section thus presents the appearance of a round disc lying in
a ring.
The head consists of two very distinct parts, a rostellum and a very swollen and
enlarged collar. No hooks could be detected either in the living or dead specimens
or in sections through the head. The rostellum is firmly imbedded in the tissue of
the intestine of the host (Fig. 15, Plate LV.). The edges of the pocket in which it
lies have grown in round the neck of the rostellum ; there seems to be no other
means of attachment. There are no hooks and the suckers are both small and weak,
and not in contact with the tissues of the host.
The collar is a large swollen structure. From the centre of its anterior face the
rostellum arises and on the same face, close to the base of the rostellum, four
minute and insignificant suckers are situated (Fig. 14, Plate LV.). In some notes
made by Dr Willey at the time of taking these parasites he states " the suckers are
not visible in the living condition." They are just visible in the spirit specimens
with a good binocular microscope under a 2 in. objective as four round spots placed
at the four ends of an equal limbed cross. In section (Fig. 15, Plate LV.) it will
be seen that the suckers are very small and unattached to the intestinal wall of the
Dog-fish.
The tissue of the collar is a spongy mesh-work which absorbs but little pigment
when stained. The nuclei of the cells lining the thin cuticle however stain, and certain
large cells in the rostellum. The collar is traversed by four bands of stout muscles
which pass from the longitudinal muscles in the body to the rostellum, where they
spread out in a brush-like manner. These four bundles of muscles doubtless permit the
movement of the body on the fixed head. Posteriorly the muscles tend to split into
eight and then into more bundles, which ultimately end in the well-marked longitudinal
muscles of the neck.
Anteriorly the body is circular in outline and the anterior end of even the mature
segments retain a circLdar outline, although the middle portion of each proglottis is
somewhat flattened. The most conspicuous feature of the anterior end of the animal
is the great development of the longitudinal muscles, these are very powerful and
regularly arranged in such a way as to recall the longitudinal muscles of an earth-worm.
The dorsal and ventral lateral canals of the water-vascular system both persist,
the ventral is however much the larger. A rather conspicuous tube which appears in
the lower edge of the posterior flap of each proglottis, which overlaps and partially
conceals the anterior half of the succeeding, almost certainly belongs to this system.
DURING HIS SOJOURN IN THE WESTERN PACIFIC.
547
ylk gid
The sections of the mature proglottides that I cut showed the reproductive organs
fairly well, the spermatozoa were already
fully formed and the vas-deferens was
in parts very much swollen with them.
The testes were large and scattered fairly
uniformly over the proglottis. Their
ducts united into two chief ducts on
each side, which ultimately unite. The
penis was not armed. The ovary lies
quite at the anterior end of the pro-
glottis, close to it was a finger-shaped
gland which I take to be the shell-gland.
The deeply staining yolk glands are un-
usually pron)inent; they are so arranged
as to form half cylinders, represented in
section by two crescents which surround
the other generative organs. The ducts
of those of each side unite and form a
wide canal which with the similar duct
from the other side of the body opens
into the duct which leads from the ovary
into the uterus.
Fig. C. Adelobothrium aetiobatidh, n. g. Diagram of
reproductive system.
(J gen. pore, male genital pore, ? qeii. pori
pore, (jlds. glands round vagina, ov. ovary
gland, t€s. testis, »(.
ylk. gld. yolk gland.
female genital
sh. gld. shell
uterus, ves. sem. vesieula seminalis.
Adelobothrium aetiohatidis, n. gen. et sp.
Head with rostellum imbedded in tissues of host but bearing no hooks. Behind
the head the neck swells out into an enormous ruff-like collar. This bears on its
anterior face four very small suckers which seem to take little or no part in the
attachment of the worm to its host. The section of the body anteriorly is circular
and so is that of the anterior end of each proglottis, but the ripe proglottides tend
to be flattened in their middle regions. Each proglottis is produced backward into a
very prominent ridge which ensheaths the succeeding proglottis to a varying extent
according to their age.
The genital pores are unilateral and irregular, but groups of three opening on
one side followed by groups of three opening on the other, succeed one another with
some regularity in certain regions.
Both dorsal and ventral, longitudinal, water-vascular canals persist, and anteriorly
the longitudinal muscles are in unusually powerful and distinct bundles.
Species. Adelobothrium aetiobatidis with the characters of the genus.
Habitat. The intestine of Aetiobatis narinari Euphrasen, amongst the folds of
the spiral valve. The host was taken at Lifu, Loyalty Islands.
w. V.
73
o48 A DESCRIPTION OF THE ENTOZOA COLLECTED BY DR WILLEY
IV. PALAIA VARAXI, u. gen. et sp.
The following account refers to a new genus of Tape-worm taken by Dr Willey
from the duodenum of a lizard, Varanus indicus. The length of the tape-worm
averaged, in the nine specimens w^hich Dr Willey obtained, about 15 — 20 mm. When
they reached me they presented the appearance roughly indicated in Fig. 16, Plate LV.
The Jiead bore an enlargement, a species of frilled or puckered turban which gave
the animal a very characteristic look and vividly recalled the figures given by E. Linton
of Thysanocephalum crispum Linton, from the Tiger-shark Galeocerdo macidatus^. For
some time, even after I had prepared and examined sections, I thought this turban
was part of the parasite. It appeared to have its origin on one side of the neck
and to grow forward and over the head. In section the tissue of this part showed
a certain resemblance to the tissue of the parasite — perhaps this was due to imperfect
preservation — and the turban showed no definite resemblance to the lining of the
alimentary canal of a Vertebrate. Neverthele.ss I gradually became con\-inced that the
turban was part of the host and not of the parasite. With great care it was possible
to dissect the head of the tape-worm out of the turban and the head showed no
trace of lesion either when viewed as a whole or when cut into sections. On opening
a specimen of Varanus indicus it was found that the lining of the duodenum was
produced into just such papillae as the turban showed, and in cutting sections of
these they were seen to histologically resemble the turban in the tape-worm. Finally
Dr Willey recollected that as the animals w^ere so deeply embedded in the tissues
of the duodenum of the Lizard that he could not remove them without breaking the
worm, he had in each case cut out a small portion of the intestinal wall to save the
tape-worm's head.
I cannot find that the worm in question belongs to any described genus and I
therefore must propose a new one, the diagnosis of which will be found on p. 550.
The name I suggest Palaia is derived from the native word for the lizard which
harbours it, and the specific name varani indicates the genus of the host.
Palaia varani has a rounded flattened head which bears no hooks (Fig. 17,
Plate LV.). It is provided with four deep but weak suckers which sink into the
neck as well as into the head. The worm retains its hold of the intestinal wall of
its host not by these suckers but by its rounded head being sunk into a pit, the
edges of the pit contract and encircle the narrow neck and the head cannot be pulled
out through this orifice mthout first slitting it, at any rate in preserved specimens.
The head as is shown in Figure 17, Plate LV. is somewhat wrinkled, and the
wrinkling is even more marked on the neck. There nre a series of longitudinal
grooves which are very deep, these give the anterior part of the body a star-shaped
section (Fig. 18, Plate LV.). When the longitudinal -wrinkles reach back to the com-
mencing transverse grooves the body becomes divided up into a number of small
quadrilateral areas. The transverse grooves which divide the proglottides are also deep
and give in section a characteristic appearance. When the proglottides however are
1 United States Commission of Fish and Fisheries. Part xvi. Report for 188B. Washington, 1892, p. 543.
DURING HIS SOJOURN IN THE WESTERN PACIFIC. 549
ripe these grooves disappear to a great extent. The width of the ripe proglottides is
about 1 mm. and in outline they are nearly square.
The reproductive opening is on one side of the body only and very irregularly
arranged, as is shown in Fig. 17 a, Plate LV. In many cases the older proglottides,
which are little more than thin sacs of ova, retain the penis in an extended condition
(Fig. 19, Plate LV.).
The whole body is covered by an unusually thick cuticle which is well shown in
Figs. 18 and 20, Plate LV. The tissue within the cuticle is rather liable to contain
lacunae or splits between the cells, and this is especially the case where a prominence
protrudes between two grooves. The most striking of the histological features is the
sheath of longitudinal muscles which separate the central part of the proglottis in
which the reproductive organs lie from the more superficial tissues. This layer consists
of numerous longitudinal bundles very definitely arranged in the manner indicated in
Fig. lb, Plate LV. They take up the pigment of certain staining fluids with great
avidity, and as the muscles .separating one proglottis from another do the same, a
very characteristic, deeply stained, ladder-like muscular system is seen in longitudinal
section. This is well shown in Fig. 21, Plate LV., which represents a longitudinal
vertical section through a P. varani near the anterior end of the animal. In the head
the longitudinal muscles diverge and break up into something not unlike a shaving-
brush.
Both the ventral and dorsal longitudinal vessels are seen in section through the
middle of the body (Fig. 18, Plate LV.). Posteriorly they tend to become smaller, and
this is especially the case with what I take to be the dorsal canal. This lies not
above but just external to the larger ventral vessel and both are included with the
tissue circumscribed by the deeply staining longitudinal muscles. The ventral vessel is
very capacious and slightly wavy in outline in that part of the body which succeeds
the head, but when the reproductive organs are well developed it diminishes much in
size. The connection of the vessels of one side with those of the other in the head
is simple and not plexiform.
The reproductive system takes up but a small part of the space of a proglottis,
that circumscribed by the above mentioned deeply stained muscles. This space relative
to the surrounding tissues however increases in size as we pass backward first to the
mature proglottides and then to those which contain the eggs in utero. The more
advanced of these is little but a thin walled sac of ova, the surrounding tissue being
to a very great extent absorbed (Fig. 20, Plate LV.). Longitudinal section through the
anterior half of the animal shows that the primordia of the generative organs do not
correspond in number or po.sition with the external annulations, they are in fact far
more numerous, but this difference is soon adjusted and by the time the generative
glands are functional each set corresponds accurately with the region between two
annulations.
Unfortunately the details of the arrangement of the testes, ovary and accessory
glands could not be made out. As was mentioned above the genital pore is unilateral,
and its distribution, on one or the other side of the body, is highly irregular. The
penis is often found in a state of protrusion and this even in the most mature
73—2
550 A DESCRIPTION OF THE ENTOZOA COLLECTED BY DR WILLEY
proglottides where the rest of the generative organs have practically disappeared (Fig. 19,
Plate LV.). The uterus is at first somewhat divided by projecting shelves of tissue,
but these become less and less prominent until in the hindermost proglottides they
have practically disappeared. The eggs are very characteristic, in the first place they
are arranged much more regularly than is usually the case, in transverse rows. They
are when isolated spherical in shape, but they show a great disposition to adhere
together in small clumps and their outline is then interfered with by mutual pressure
(Figs. 19 and 20, Plate LV.). These clumps perhaps most usually consist of two eggs,
but three pre.s.sed together or even four are often met with. The egg shell is single
and the nucleus large, the average diameter of the whole egg is about "025 mm.
The systematic position of P. varani is near that of the Dr Max Liihe's recently
described Oochoristica'^, but I think sufficient differences exist to justify the establish-
ment of a new genus. Until we have a fuller account of the anatomy of Oochoristica
it will be impossible to determine the exact relationship of the two forms.
The following are the features of the proposed new genus Palaia : —
Palaia varani, n. gen. et sp.
The rostrum is absent and no hooks are present. Four weak suckers are found
on the head. Immediately behind the head is a narrow constriction or neck. The
head is deeply embedded in a pit in the substance of the alimentary canal wall of
the host. The edges of the pit have narrowed and by means of the button-shaped
head the parasite is kept in po.sition. The body is deeply grooved with longitudinal
furrows which with the transverse furrows form small quadrilateral areas. Genital pore
unilateral and very irregularly arranged. The longitudinal muscles surrounding the
central part of each proglottis, in which the generative glands lie, ver}' well marked
and divided into very definite bundles. Ova "025 mm. in diameter, more or less
regularly arranged, sometimes aggregated together in clumps of two, three or four.
Species. Palaia varani, with the characters of the genus.
Habitat. The duodenum of the lizard Varanus indicus.
V. PHYLLOBOTHRIUM DIPSADOMORPHI, n. sp.
In von Linstow's " Compendium der Helminthologie " 1S78, and its " Nachtrag "
1889, some eighty-six species of snake are mentioned infested with entozoa, but the
number of species of adult Cestode catalogued amounts to but thirteen belonging to
six genera. In the British Museum Catalogue of Snakes, Bou longer describes over
1600 species of Snake, and it is thus evident that at the present time but a small
percentage of the Ophidia have been searched for parasites and that those which
have been investigated contain a comparatively small number of genera of Tape-worm.
The species I am now about to describe belongs undoubtedly to the genus Phyllo-
buthrium and was taken from the intestine of a " Malagea " snake Dipsadoviorphus
1 Zool. Anz. XXI. 1898, p. 6.50.
DURING HIS SOJOURN IN THE WESTERN PACIFIC. 551
irregularis (Merrem.) at Karavia at the head of Blanche Bay, New Britain, on the
18th of July, 1897. Dr Willey distinctly recollects taking tape-worms from a "Malagea"
and his Journal contains the following entry: July 18th, 1897. "In the afternoon,
dissected a large lizard, 'a palai,' with groups of yellow spotted scales, and a large
snake, 'a malagea,' nearly 42 inches loug....Acarines, Cestodes and Nematodes and
Pentastomids." There is thus every reason to believe the label in which the above
statements are recorded is correct. The reason why I am so precise in this affair
is that hitherto the genus Phyllobothrium has only been recorded from Fish, and its
presence in a Snake demands some explanation. Unfortunately I can offer none, the
only excuse for the snake being that it was living in sight of the sea.
Van Beneden described in 1850' two species of Phyllobothrium, Ph. lactuca in
Mustelus vulgaris and Ph. tridax in Squatina angelus. No trace of a small sucker
in each bothria appears in his figures. In 18.58 he mentions as characteristic of the
genus a sucker near the external edge of the bothria. In this paper- he describes Ph.
auricula from Trigon pastinaca.
In 18.55 WedP described another species, Ph. gracile, taken from the intestine of
Torpedo marmorata, here again the suckers are not evident.
In 1866-67 P. Olsson-* described a new species. Ph. acanthiae vulgaris, taken
from the intestine of Acanthias vulgaris and some undetermined species from Clupea
harengus and Lahrus maculatus'.
In 1870 van Beneden in his " Les Poissons des Cotes de Belgique"" mentions
Ph. brassica from Spiiuw acanthias and Ph. fallax fi'om Raia rubus.
The longer of the two specimens at my disposal is 7 cms. in length, the width
1'.5 nmi. The latter does not vary much, the anterior segments except those immediately
behind the head being almost as broad as the posterior. Tlje hindermost segments
are very elongated, some attaining a length of 3 or even 4 mm. The constrictions
between the segments are slight and there is no overlapping of a proglottis by its
predecessor.
The head bears four unstalked lappets and thus to some extent resemble a four-
fold clover leaf, as each lappet is indented on its outer edge and somewhat heart-
shaped (Fig. 22, Plate LV.). Round the outer border a well marked rim extends,
which from time to time gives off a slight ridge which runs for a little distance
towards the apex of the heart-shaped lappet and then fades away.
I could discover no trace of any suckers, but as I could not sacrifice either of
the two heads to the microtome I cannot speak quite positively upon this point.
Suckers however were not observed by the earlier investigators of the genus.
' "Recherches sur la Faune littorale de Belgique. Les Vers Cestoides. " .l/rai. Ac. Helgique, xxv. 18.50,
p. 120.
- Mimoire iur les Vers Intestinaux, Paris, 18.58,
» SB. Ak. Wien, xvi. 1855, p. 373.
* Acta Univ. Lund. in. 1866—67.
' loc. cit. IT. 1867.
" Mem. Ac. Belgique, xxxviii. 1877, pp. 10 aud 17.
552 A DESCRIPTION OF THE ENTOZOA COLLECTED BY DR WILLEY
One of my specimens head the lappets folded one on another so that the head
appeared spherical and resembled the Fh. lactuca of van Beneden, the other fortunately
had its lappets spread oiit as shown in the figure.
Zschokke' has given an excellent account of the anatomy of a proglottis of this
genus and I have little to add to what he has written. I did not observe any
transverse vessels uniting the longitudinal canals of the water-vascular system, but
the latter give off numerous fine branches. The uterus as it first appears is a rod-
like body, its walls then increase laterally b)' numerous pockets. These pockets are
separated from one another by dissepiments which end in a trifid or quadrifid edge.
In the oldest proglottides which were at my disposal the uterus has ruptured
about the centre of the dorsal surfice and I am inclined to think that this normally
takes place, for even in the segments in which the uterus is not full of ova there
is a marked thinning of the wall of the uterus and body at this spot.
The following are the characters of the species : —
Phyllobothrium dipsadomorphi, n. sp.
Head with four flattened heart-shaped lappets, which bear no sucker (?). A well-
marked raised rim which gives off a few radial ridges surrounds the lappets. These
pass a little way towards the apex of the lappet. Breadth of body very uniform.
Genital pores unilateral and irregular, two or three to the left and then two or three
to the right.
Habitat. Intestine of Dipsadomorphus irregularis, Blanche Bay, New Britain.
VI. COELODELA KUVARIA, n. gen. et sp.
The following is an account of a Tape-worm taken from the intestine of a fi-uit-
eating Pigeon, Carpophaga van-ivycki, shot near Karavia in the Gazelle Peninsula,
New Britain. It will be seen from the sequel that I have been obliged to pro-
visionally propose a new genus for this species. It is always a somewhat risky thing
to establish a genus on a single specimen, and it is especially so when that genus
has to be characterised by structural peculiarities, the very discovery of which has
entailed the destruction of the specimen as a whole. The differences between this
specimen and others with double sexual organs and pores will, I think, justify the
establishment of a new genus for which I suggest the name of Goelodela. The generic
name has reference to the extraordinary development of the spaces of the water-
vascular system. The specific name Kuvaria is derived from the native word Kuvar,
which denotes a pigeon. From the fact that this animal inhabits the pigeon, a bird
extremely liable to be shot, it is to be hoped that it will not be long before moi-e
examples of this parasite are available for research.
The length of the worm is some .50 mm., its greatest breadth .5 mm., the breadth
of its head 1 mm. The ventral surface of the proglottides is slightly hollowed and
' Mem. Inst. Geiiev. xvii. 1886—1889, p. 317.
DUEING HIS SOJOURN IN THE WESTERN PACIFIC. 553
the dorsal surface correspondingly arched. The genital pore is conspicuous on each
side of each proglottis.
In examining the head of this tape-worm, soaked in oil of cloves to render it
transparent, it became evident that there were no hooks. The sections through the
head confirmed this observation. Not only are hooks absent but practically there is
no rostellum. A small portion — but a very small one — of the head lies in front of
the anterior edge of the suckers. The latter are deep and somewhat retracted in the
specimen and their hollow, when looked into, showed a fold or valve which has been
drawn in Fig. 23, Plate LV. Two suckers are dorsal and two are ventral. Each
sucker has a number of longitudinal muscles attached round its edge which run
down the neck and which contracting together would serve to withdraw the whole
sucker or contracting alternately would rotate it about its centre.
Inside the head the parenchyma surrounds a nervous ganglion situated between
the suckers, this histological state of the material does not allow one to state more than
that ganglion cells and fibres are to be made out in this mass. Laterally the ganglion
gives off two nerves which run down the animal just external to the lateral vessels.
The extreme anterior end of the head is occupied by a close plexus of fine
excretory vessels which splitting into two plexuses passed one on each side of the
body between the right and left suckers of each side (Fig. 23 a, Plate LVI.). These
plexuses consist of some 10 or 12 vessels at any one level, splitting into two and
running into one another. The diameter is about that of the dorsal longitudinal
vessel and far smaller than that of the ventral lateral vessel. Shortly behind the
level of the suckers, the plexuses gather themselves up into a very fine dorsal vessel
and a much larger ventral vessel. The latter communicates with its fellow by a trans-
verse vessel at the posterior end of each segment, the former gives off a number of
small diverticula which appear to end blindly but probably open into minuter canals.
The most striking anatomical feature of this tape-worm is the spacious nature
of tlie canals of the water-vascular system. The dorsal vessel soon disappears but
the ventral vessel maintains a capacious cavity which communicates with its fellow
at the posterior border of each proglottis (Fig. 23 a, Plate LVI.). At the point of
emergence of the transverse vessel from the longitudinal the latter slightly contracts
and the entrance to the former is guarded by a valve which only permits fluid to
enter them when flowing from before backwai'ds (Fig. 24, Plate LVI.). The figure
just mentioned hardly does justice to the large space of the proglottis which is taken
up by the transverse vessel, and this space relatively increases as we pass from
before backward. Fig. 25, Plate LVI., is a transverse section, cut slightly obliquely
through one of the most posterior proglottides. On the right side the spacious uterus
containing ova is seen and on the left the still more spacious transverse commissure
of the water-vascular system. It will be noticed that the latter has an even greater
dorso-ventral width than the uterus. The same appears true of the anterior-posterior
diameter and the Figure 26, Plate LVI., shows that the lumen of the transverse
water-vascular canal is as great if not greater than that of the uterus and occupies
at least half the cubic content of each proglottis. I am unacquainted with any other
species of tape-worm which has so large an extension of its excretory-system.
554 A DESCRIPTION OF THE ENTOZOA COLLECTED BY DR WILLEY
The most posterior twenty segments or so of this worm were removed and divided
into three portions. Each of these was cut into sections in a different plane. The
segments at this end were, as was to be expected, mature and the uterus was spacious
and crowded with ova in well developed spherical egg-shells. The uterus is single and
common to the two sets of generative organs as in Dipylidium and Ctenotaenia.
When the ova first pass into the uterus it is a branching sac, the numerous
diverticula of which are separated by well developed and bulky masses of parenchyma,
but as the segments age this parenchyma tends to be absorbed, the partitions it
creates between the diverticula of the uterus disappear until in the last stages the
uterus is a cavity with practically no subdivisions, an occasional slight strand projecting
into its lumen being the only trace of its former irregular shape.
At this most ancient and posterior end of the animal the enormous development
of the transverse canal connecting the two ventral lateral vessels is fully apparent.
They have attained so large a size as to equal that of the uterus and the two
together form very nearly the whole of each segment, for the walls of both have
become very thin. No trace of the small dorsal lateral vessel exists at this end of
the animal. At the junction of each transverse vessel with the ventral lateral there
is a well developed valve which permits a flow of the fluid contained therein only
in the direction from before backward. The valves are very regularly arranged, as is
seen in Fig. 24. The only other structures to be found in these posterior segments
are the two nerve cords, each very flattened and closely applied to the outer surface
of the dorsal lateral canal, and the remains of the genital ducts with their openings
one on each side of each segment. The tissues of the body are at this stage very
much reduced and the parenchyma is largely absorbed.
The determination of the .systematic position of this worm is a matter of great
difficulty. In the last few years a considerable number of new genera have been
established for the tape-worms of Birds. Some of these like the species in question have
double genital pores on each proglottis. Amongst these may be mentioned the Cotugnia
and Amahilia of Diamare', and the Diploposthe of A. Jacobi-. The specimen Dr Willey
has brought home differs from these species in the entire absence of hooks. Both
when examined in oil of cloves as a whole and when cut into sections and examined
in detail, no trace of a hook was to be seen. However too much reliance must not
be placed on this point, for as Professor Railliet has been kind enough to remind
me the hooks of the tape-worms of birds are very apt to fall off and are not
unfrequently left behind in the host.
The rostellum of my form is so rudimentary that it can hardly be said to exist
at all. Another feature in which this species differ from — at anj- rate — Coturjnia is
that the uterus in the older proglottides is a spacious, definitely- walled chamber with
nothing but ova in its cavity whereas in the last named species it is " representato
da cellette riempite da una massa parenchimatosa in cui si trovano le nova." The
' Ball. Sue. Napoli, Ser. 1, vii. 1893, p. 9, and Gentrhl. Bakter. xxi. 18'.i7, p. 862 and xxv. 1899, p. 35-2 ;
see also Cohn, Zool. Anz. xxi. 1898, p. 557.
■•^ Zool. Jahrb. Anat. x. 1897, p. 287, and CentrhK Bakter. xxi. 1897, p. 873. The question as to the generic
identity or distinctness of tlie last two forms mentioned is discussed in the memoirs here cited.
CUBING HIS SOJOURN IN THE WESTERN PACIFIC.
555
9 gen pore.
■■3 gen pore.
Fig. D. Diagram of the uterus and male reproductive organs of
Coelodehi kiwaria.
J and (f gen. pore, male and female genital apertures, glds.
round vagina, I. v. lateral water-vascular vessel, tes. testis, ut. uterus,
V. valve in l.v., ves.sem. vesicula semiualis.
arrangement of the generative organs with the compact female glands and the testes
scattered on the dorsal side of each proglottis is also unlike what occurs in the above-
mentioned forms.
In some respects the an-angement of the genitalia recalls that which obtains in
the genus Muniezia but the
uterus is single. The first trace
of the reproductive organs ap-
pears in the younger segments
shortly behind the head, the
first structure to appear and
the la.st to disappear when the
proglottides are full of ova is
the cirrhus and its base. The
cirrhus bulb is large, when the
cirrhus is protruded it leaves
the proglottis about the junction
of the anterior two-thirds with
the posterior third. Close to it
opens the vagina. The vas de-
ferens leaves the cirrhus bulb
coils slightly and enlarges into
a large vesicula seminalis, which
even in the proglottides of the posterior end contains bundles of spermatozoa. The vas
deferens is then continued on and branches into a number of secondary ducts which end
in the dorsally arranged testes. There is no sign that the testes are double but probably
they are so. That is, probably no single testis lobe opens into a vas deferens which
runs both right and left. Each lobe probably opens on to either the right or the
left side of the animal. With the exception of the uterus the female organs are
also paired, and open on each side of each proglottis. The ovary and shell-gland and
vitellaria seem to be all aggregated near one edge of the animal, though the
histological state of preservation of the material did not allow this point to appear
with absolute certainty. The uterus is not paired. The first sign of it is a transverse
band of deeply staining cells which becomes differentiated right across the segment in
the centre of the body but nearer the anterior than the postei'ior limit of each
proglottis. A few segments behind that in which this transverse differentiation occurs
it is seen that further differentiations arise at right angles to the first and continuous
with it. Thus in this stage the uterus is solid and resembles the head of a rake,
with all the teeth pointing posteriorly. A little later a lumen arises within these
strands which is at first lined by a well marked epithelium. As the proglottis ages
the number of diverticula increase and their lumen swells, the intervening parenchyma
being at the same time absorbed. Thus it comes about that at first a lobed uterus
arises and then by the disappearance of the dissepiments the uterus becomes one
single undivided, smooth walled receptacle full of eggs. The ova are spherical with
one or two deeply staining yolk granules and an egg-shell which stands off from
w. V. 74
556 A DESCRIPTION OF THE ENTOZOA COLLECTED BY DR WILLEY
the body of the egg and is in the preserved specimens in all cases crumpled and
distorted.
The above-mentioned facts show that the tape-worm in question differs markedly
from the other genera with double genital orifices, which are found in birds. To
bring this out more clearly I have prepared the following abstract of the more
important anatomical features in which Cotugnia, Amabilia and Diploposthe differ from
the one in question.
Cotugnia. Rostrum rudimentary with a great number of very minute hooks.
Suckers large. Uterus full of a parenchymatous mass in which the eggs are found
in the mature proglottides. Water-vascular system not particularly large. Habitat,
intestine of certain Birds.
Amabilia. The most important feature of this genus is that the cirrhus is double
but the female generative organs are single and in the median line. The male organs
are lateral and those of one side are distinct from those of the other (Cohn). The
vagina is median. Habitat, intestine of Phaenicopterus roseus (antiquorum).
Diploposthe. The ovary is paired but lies in the middle line, a vagina passes
off from each and opens laterally close to the cirrhus. The testes are aggregated near
the centre of each segment. The lumen of the uterus in the ripe proglottis is spacious
and not at all occluded by parenchyma. Habitat, small intestine of Fuligida ferina
and other ducks.
I should put together the following as the features upon which, provisionally,
this genus is founded : —
Coelodela Kuvaria n. gen. et sp.
Rostrum rudimentary bearing no hooks. The male and female organs are double
vnth the single exception of the uterus. The vas deferens and the vagina open on
each side of each proglottis. In mature specimens the uterus is spacious -with practically
smooth walls and the lumen is not divided up by dissepiments. The water-vascular
spaces attain very large dimensions and this is especiall)' true of the transverse
canals. The dorsal vessel does not reach beyond in the neck.
The species has the characters of the genus.
Habitat. The intestine of a fruit-eating Pigeon. Carpophaga van wycki shot at
Karavia, Gazelle Peninsula, New Guinea.
It is worthy of notice that the only other tape-worm recorded from this genus
of fruit eating Pigeons is the Taenia insignis of F. Steudener', who found the parasite
in the alimentary canal of Carpophaga oceanica Lesson, a bird which is recorded in
the British Museum Catalogue under the name Globicera pacifica. The anatomy of
Steudener's T. insignis shows little or no resemblance to that of the form described
above.
' Abh. Ges. Halle, xiii. 1877, p. 277.
DURING HIS SOJOURN IN THE WESTERN PACIFIC. 557
VII. PROSTHEGOGOTYLE DIOMEDEAE Fuhrmann, n. sp.
The following is a list of the Cestoda known to inhabit the Albatros :
1. Prosthecocotyle^ sidciceps Baird. This species was described by Baird in 1859-
from the intestines of Diomedea exulans L. and again by von Linstow in his report'
on the " Challenger " Entozoa, under the name Taenia diomedeue though he suggests
tliat it may be the same species as Baird's. Von Linstow's specimens were found
in the stomach of D. albatrus Pall, living in the Pacific.
2. P. torulosa Linstow, was described by von Linstow under the name Tetra-
hothrium torulosum, it also occurs in the stomach of the D. albatras from the North
Pacific.
.3. P. nmhrella Fuhrmann. This species is described from an unidentified species
of Diomedea taken in an unknown locality.
Amongst the tape-worms brought back by Dr Willey were some specimens taken
from the duodenum of Diomedea exulans L. which very obviously belong to the
Monticelli's genus Prosthecocotyle. Dr Fuhrmann of Neuchatel who is monographing
the tape-worms of birds and who has paid very esj)ecial attention to the genus in
question pronounces Ur Willey 's specimens to form a new species which he characterises
as follows :
Prosthecocotyle Diomedeae Fuhrmann, n. sp.
"The length of the Tape-worm is 140 mm.; the maximum width 1mm.
Segments very short. Scolex unarmed, with O'o mm. length 0-4:3 mm. The shape
of the head is quadrangular. The suckers are oval with very powerful ear-shaped
projections. The external longitudinal muscles form bundles of 3 — .5 fibres, the
internal of 3.5 — 40 fibres. The genital cloaca is very muscular. The vagina opens
on a prominent papilla, on the dorsal side of which is the opening of the cirrus.
The receptaculnm seminis has a very strong muscular sheath and is situated on
the outer side of the water-vascular system. The spherical cirrus pouch has a
diameter of 0'086 mm. The testes are 17 — 20 in number. Eggs?"
Habitat. Host. Diomedea exulans L., in duodenum. Locality. Western Pacific.
I here append a-s a supplement to Dr Fuhrmann's diagnosis a few notes I have
made on the specimens of this new species.
At the broadest part of the body which is some little way in front of the
posterior end the width of the proglottides is about one millimetre. The antero-posterior
width of a proglottis is here about one-tenth of a millimetre. In front of this
' Monticelli, F. S., Boll. Mus. Torino, vii. 1892, p. 8. Lonnberg, Hamburger Magalhaenische Sammelreise,
1896, p. 9. Fuhrmann, 0., Zool. Anz. xxi. 1898, p. 385, ibid. xxii. 1899, p. 180, and Centrbl. Bakter. xxv.
1899, p. 863.
' P. Zool. Hoc. London, Part xxvii. 1859, p. 111.
' Challenger Reports, xxiii. Part lxxi.
74—2
558 A DESCRIPTION OF THE ENTOZOA COLLECTED BY DR WILLEY
region this dimension decreases towards the head (Fig. 28, Plate LVI.). Towards the
tail it increases and the increase is accompanied by a diminution of the lateral diameter
until the breaking off proglottides are almost as long as they are broad.
Looking down on the head the anterior surface presents two funnel-like depressions
which are lateral]}- placed. The lips of the funnel — the " ears" of FuhiTuann — are not
complete but a slit exists laterally. The edges of the slit usually overlap, they run
into one another at an acute angle situated laterally between the dorsal and ventral
suckers and about equidistant between the anterior and the posterior limits of the
head (Fig. 27, Plate LVI.). The whole relation of the edges of the funnel resemble
those of the spathe of an Arum. Had the edges fused instead of overlapping the
funnels would then have been complete and would have resembled those described
by Perrier' in his genus Duthiersia. In this genus the funnel opens by a small
pore at the narrow end and the edges are much folded and crinkled.
The two dorsal and the two ventral suckers are completely separated from one
another anteriorly by the funnel and posteriorly by a slight groove or line which
runs a little distance backwards from the posterior limit of the funnel. The right
and left funnels, both dorsal and ventral, tend however to fuse and the median dorsal
and ventral ridge which lies between the right and left sucker of both aspects
diminishes anteriorly until the suckers tend to run into one another. Thus the outline
of the rim of these suckers resembles a Greek &> with the upper limbs joined together.
The first proglottides are narrow and well marked, posteriorly however they swell
out and their anterior and posterior edges overlap the narrow stalk which connects one.
It is very difficult to explain the exact relationship of the flaps of the funnels
to the suckers, originally it looks as if the outside of the flaps on. each side were
the original suckers and that the ridge which has been described above is a later
development. In transverse section it is seen that this ridge is separated from the
surface of the flap by a basement membrane, and it has the appearance of being
stuck on (Fig. 29, Plate LVI.). The four longitudinal vessels appear in the same figure,
they continue as far forward as the second section in one series I had cut and then
simply run one into another, there is no such plexus in the head as there is for
instance in C. kuvaria. All through the head the diameter of the vessels remains
constant, and the dorsal vessels are slightly smaller than the ventral.
The reproductive pore is in every proglottis laterally placed and on the left.
Besides the specimens just described I have in my possession from the same host
several long fragments of a tape-worm which I am unable to identify. They were given
to me by Mr F. F. Laidlaw, of Trinity College, who had extracted them from an
Albatros, but the want of preservation and the cold storage to which the bird had
been subjected on its journey to England had apparently injured the specimens and the
state of their preservation was bad. Several of the fragments attained a length of
6 or 7 in. but they were very fragile and easily broken. Although I made a most
careful search I was unable to discover a single specimen with a head.
1 Arch. Zool. (jp. II. 1873, p. 349.
DURING HIS SOJOURN IN THE WESTERN PACIFIC. 559
III. NEMATODA.
The collection of Nematodes made in the East by Dr Willey comprised four
genera and six species, one of the latter new. I am much indebted to Professor
M. Stossich of Trieste who named the specimens for me, and has furnished me with
a copy of his diagnosis of the new species which I append (pp. 560, 561).
I. PHYSALOPTERA OBTUSISSIMA Molin.
Molin. SB. Ak. Wien, xxxix. 1860, p. 654.
Drasche. Verb. Ges. Wien, xxxii. 1882, p. 127.
Stossich. Boll. Soc. Adriat. xi. 1889.
This species is recorded from the stomach and intestine of the following snakes :
Erythrolaviprus aesculapi, L. {Coluber); Spilotes pullatus, L. (Coluber); Xenodon severus,
L. (Coluber) ; Lachesis lanceolatus, Lacep. (Coluber). The remaining names of hosts are
Fitzinger's and cannot be identified : Ophis coeruleus, Ophis rhodogaster, Ophis treuen-
steinii, Pseudophis cinerascens, Cloelia plumbea, Cloelia fasciata, Lygophis regius.
My specimens were taken from the intestine of the snake Dipsadomorphus irregu-
laris (Merrem) found by Dr Willey in New Britain. As all the snakes from which
this species of Nematode has hitherto been taken are South American this discovery
materially increases the geographical range of the parasite.
II. PHYSALOPTERA RETUSA Rud.
Spiroptera retusa, Dujardin. Hist. nat. d. Helm. 1845, p. 104.
Diesing. Syst. Helminth, ii. 1851, p. 236.
Molin. SB. Ak. Wien, xxxix. 1860, p. 652.
Schneider. Monogr. d. Nematod. 1866, p. 65.
Stossich. Boll. Soc. Adriat. XI. 1889.
This species has hitherto been recorded from the oesophagus, stomach and intestine
of the following Lacertilians : Tupinambis teguiooin, L. (Lacerta) ; Tupinambis nigro-
punctntus, Spix ; Ameiva snrinamensis, Laur. (Seps) ; Scleroporus undulatus, Merr.
(Uromastyx) ; Amphisbaena alba, L. ; Ophiodes striatus, Spix. (Pygopus). The Podinema
scripta and Euprepis Spixii of Fitzinger, also mentioned as hosts of Ph. retusa, are
nomina nuda.
The above are all Lacertilians and all from Brazil. My specimens were taken
fi-om the intestine of the "malagea" snake Dipsadomorphus irregularis (Merrem) taken
in New Britain. Thus the zoological and geographical distribution of the parasite is
widely increased.
560
A DESCRIPTION OF THE ENTOZOA COLLECTED BY DR WILLEY
III. PHYSALOPTERA VARANI Par.
Parona. Ann. Mus. Genova. Ser. 2, vii. (xxvii.), 1890.
Parona's specimens came from the stomach of a Varamis
bengalensis, Daud. (Tupinamhis), taken at Palon, Pegu. Dr Willey's
occurred in the stomach of Varanus indicus, Daud. {Tupinamhis),
taken in the Western Pacific Isles.
IV. ECHIN0CEPHALU8 STRIATUM Montic.
Monticelli. Boll. Soc. Napoli, Ser. 2, ill. Anno III. 1889.
Monticelli's specimens on which the species is founded were
taken from the stomach of a dog-fish, Sct/llium sp. captured off
Payta (Peru). Dr Willey's specimens occurred in the intestines of
the four-spined Sting Ray, Aetiohatis narinari Euphrasen, taken
at Lifu, Sandal Bay, Loyalty Islands.
Fig. E. Kchiitucephalm
striatus Monticelli ?
Anterior end of the body.
V. SCLEROSTOMUM APPENDIGULATUM Molin.
Kalicephalus appendicidutus Molin. II sottordine degli Acrofalli
IX. 1861.
Mem. 1st. Veneto.
Stossich, M. Strongylidae, Boll.
Soc. Adriat. xix. 1899.
Dr Willey found his specimens in
the oesophagus of the snake Dipsadomor-
phus irregularis Merrem, the " malagea "
of the Malays. It has previously been
recorded from the intestine of Xenodon
merremi, Spix (Ophis), Xenodon sever us,
L. (Coluber) ; Leptophis liocercus, Wied.
{Leptophis abaetulla) ; Spilotes pullatus,
L. ; Drymohius bifossatus, Raddi {Coluber
lichtensteinii); Drymohius bivittatus, Dum.
and Bibr. (Leptophis) 1 Rhadinaea mer-
remii, Wied. ; and one or two others
such as Ophis coeruleus and Cloelia
plunibea which cannot now be identified.
The previous hosts however are South
American and thus Dr Willey's find has
widened the geographical distribution of
the parasite.
VI. GNATHOSTOMA SHIPLEYI Stossich.
This new species of GmttJiostoma was taken from the duodenum of the Albatros,
Diomedea exulans L. Hitherto the members of this g-enus have not been recorded
from any bird. I append the description of the new species kindly furnished me by
Professor M. Stossich'.
I Boll. Soc. .-idriat. xx. 1900.
Fig, F. Sclerostomum appendiculatuiii Molin.
1. Anterior end of the body seen dursally,
2. The same seen laterally,
3. Genital bursa of male seen veutrally.
DURING HIS SOJOURN IN THE WESTERN PACIFIC.
561
Per gentilezza del Signer A. E. Shipley di Cambridge (Inghilterra) ebbi a dispo-
sizione tre nematodi raccolti dal Dott. Willey nel duodeno di una Dioviedea exulans
{Western Pacific). Sono un ma.schio e due femmine appartenenti all' interessantissimo
genere Gnathostovia Owen (Cheiracanthus Diesing), genere questo rappresentato fino ad
ora da un numero molto limitato di specie
viventi tutte nel tubo digerente dei verte-
brati, fixtta eccezione per gli uccelli. Le
specie descritte sono : pei mammiferi la
G. spimgerHin Owen (Felis tigris, concolor,
catus e Putorius vulgaris), la G. hispidum
Fedtsch. (Sus scrofa fer. et doin. e Bos
taurus), la G. siamense Lev. {Homo sapiens)
e la G. socialis Leidy (Mustela vison) ; pei
rettili la G. horrida Leidy {Alligator mis-
sissipiensis) e per i pesci la G. gracilis
Diesing ( Vastres cuvieri e Sudis gigas) ;
cosicche tanto piii interessante riesce la
scoperta del Dott. Willey, inquantoche la
G. shipleyi rappresenterebbe la prima ed
unica specie di questo genere vivente negli
uccelli.
Presenta il corpo lungo cilindrico, con
la cute grossa provveduta di una distinta
striatura trasversale e di una fittissima
striatura longitudinale. II corpo anterior-
mente si assottiglia di molto, la cute invece
si solleva (circa come nelle Physaloptere) e
forma una specie di testa di un' eleganza
sorprendente. La bocca e terminale e
limitata da due labbra dorsoventrali che
appariscono trilobate, e ogni lobo all' estre-
mita con una minutissima papilla ; le labbra
sostengono due grandi disclii cefalici in
posizione pure dorsoventrale, larghi, pos-
teriormente bilobi e col margine armato di
forti aculei, i quali verso 1' estremita dei
lobi aumentano gradatamente in grandezza ;
questi due dischi oefalici sostengono la cute sollevata del capo, c dall' osservazione i'atta
mi sembra che debbano essere suscettibili a movimenti dal basso all' alto. Poco sotto
questi dischi cefalici si vedono due papille cervicali (?), una dorsale ed una ventrale,
tricuspidate, di sviluppo imponente e molto sporgenti ; da queste papille cervicali prin-
cipiaiio le lamcllo, che in questa specie sono tutte semplici e disposte in sole quattro serie
longitudiuali, due dorsali e due ventrali, che si estendono circa fino alia meta del corpo.
Fig. G. Gnathostoma shipleyi Stossieh.
1. Lateral view ot anterior end of boLly, 2. Dorsal view
of the same, 3. Posterior end of the female, 4. Posterior
end of the male, 5. Egg.
562 A DESCRIPTION OF THE ENTOZOA COLLECTED BY DR WILLEY
L' estremita caudale del maschio si presenta attortigliata a spira, con 1' apice
arrotondato e cou una borsa genitale pochissirao sviluppata ; le papille caudali deli-
catissime e di difficile osservazione, dovrebbero essere in numero di 17, quattro paia
preanali e quattro paia postauali disposte in due gruppi, piu una papilla terminale
air apice caudale con 1' estremita multicuspidata ; cirro destro molto robusto e di uno
sviluppo del tutto speciale. L' estremita caudale della femmina e parimente conica con
r apice arrotondato e sotto la cute si osservano tre piccolo papille, una apicale e due
laterali. La vulva si apre poco sopra la meta del corpo e le uova numerosissime,
contenenti 1' embrione, sono di forma ellittica a guscio grosso e liscio.
Lunghezza del maschio 15™".
Lunghezza della femmina 35™".
IV. NEMATOMOEPHA.
The solitary Gordian worm brought back by Dr Willey turns out to be a new
species. Dr L. Camerano' of Turin has kindly described the worm for me and I
quote his description.
The worm was found amongst the roots of certain water weeds in the muddy
banks of a stream at the elevation of several hundred feet at the head of Blanche
Bay, New Britain. It was at this place that Dr Willey first took Peripatus novae-
hrittaniae. Judging from its forked tail it is a male.
There is no evidence as to the animal in which the Gordian worm may have
been parasitic.
I. GORDIUS WILLEYI Camerano.
"Un esemplare J della Nuova Britannia raccolto dal dottor A. Willey nel 1897 —
Nuovo Museo di Cambridge (Inghilterra).
" Lunghezza m. 0,27. — Larghezza massima m. 0,0008.
" La parte anteriore del corpo e alquanto appunlita : Testremita posteriore ha i
lobi luughi poco piu di mezzo millimetro alquanto divergenti, ingrossati e rotondeg-
gianti alia loro estremita (Fig. .30, Plate LVI.). La lamina postcloacale e ben spiccata, a
ferro di cavallo.
" Lo strato cuticolare esterno e privo di areole papillari e di produzioni peliformi :
presenta le linee incrociate solite a trovarsi in tutte le specie del genere Gordius
come io I'ho inteso nella mia Monografia dei Gordii.
" La colorazione generale e di color bruno, senza alcuna sorta di macchiette chiare :
la calotta anteriore e bianca : il collare nero ben spiccato ed esteso per poco meno
di mezzo millimetro : le striscie brune laterali poco spiccate.
" Questa specie appartiene a quella sezione del genere Gordius, in cui lo strato
cuticolare esterno non ha speciali formazioni rifrangenti che vengono ad affiorare alia
■ Atti Ace. Torino, xxxiv. 1899.
DURING HIS SOJOURN IN THE WESTERN PACIFIC. 563
Superficie esterna dello strato cuticolare stesso, e rientra nel sottogruppo da me stabilito
nella monografia sopra citata, caratterizzato dalla presenza di un uollare nero ben
spiccato. Questo gruppo comprende appunto le specie fino ad ora state deseritte delle
Indie Orientali come il G. fulgur Baird, il G. doriae Camerano, il G. horsti Camer.,
il G. salvadorii Camer.
" II Gordius willeyi si differenzia facilmente da queste specie per la forma dei
lobi deir estremita posteriore ; facilmente pure si distingue dal G. villoti Rosa, pei
caratteri della cuticola esterna e per ([uelli dei lobi posteriori."
V. LINGUATULIDAE.
I. POROCEPHALUS TORTUS Shipley.
In an article entitled "An Attempt to Revise the Family Linguatulidae'" published
in 1898, I gave a short description of a new species of this family of parasites which
I named Porocephcdus tortus. The specimens of this animal had been sent home by
Dr Willey and the description was written before he had completely sorted the
valuable material he had accumulated in the far East. He has now found some more
specimens and two more stages in the development of the animal, and has further
given me some more details as to the host and the dimensions of the parasites when
in the fresh state. Before mentioning these, however, I insert the description of the
species made from spirit specimens, quoted from the above-mentioned article.
" The young form is very distinctly ringed, the number of rings is about 2.5.
In the mature forms the development of the genital organs has so stretched the
skin that the rings have disappeared, at the same time the body has become curiously
twisted as is shown in the figure, and this characteristic feature has suggested the
specific name which I have given to this form. The cephalothorax which includes
the first 3 or 4 rings is well marked otf from the bod}'. The rings, as the body
stretches, disappear last from the region behind the well-marked neck.
"The hooks are single, the inner pair slightly in advance of the outer. The mouth
almost oval, and a little posterior to the inner hooks, a chitinous half-cylinder runs
back from it along the posterior face of the oesophagus.
" Length of mature female 40 mm. The longest specimen measured 3 mm.
more but the others were very constant in their length. The head measured 5 mm.
The width of the body which was very constant behind the neck, only slightly
tapering towards the tail is 2, .5-8 mm. The length of the immature female whose
rings are just disappearing is 35 mm., and the length of the young form with well-
marked rings is 10 mm., its breixdth about 1 mm.
"Habitat. In the body of Dipsadomorphus irrerjularis, from New-Britain. Found
by Dr Willey."
The parasites were taken from within the body of a snake — Dipsadomorphus
irregularis Merrem, the Malagea of the natives, but I had not at the time I wrote
1 Arch, parasit. i. 1898, p. .52.
w. V. 75
564 A DESCRIPTION OF THE ENTOZOA COLLECTED BY DR WILLEY
the above diagnosis appreciated the fact that this snake was discovered inside the
stomach of another called by the natives a Taluluka and kindly identified by
Mr Bouleuger as Nardoa boa F. Miill. In the British Museum catalogue this last-
named snake is recorded from the Duke of York Island and New Ireland; as Dr Willey's
specimen was taken inland near Blanche Bay, ^ew Britain, its range is slightly
extended.
The Dipsadomorphus has suffered to some extent for the digestive action of the
Nai-doa. The flesh on the skull was beginning to disappear but the part which had
suffered most began about six inches behind the head and extended some six or
eight inches. Here the ventral body wall of the body was completely digested away,
leaving the ribs sticking out of the sides of the body like rows of tomb-stones. It
was within the cavity thus laid open that Dr Willey found the parasites which he
describes as " devouring " the swallowed snake.
Which of these snakes is the host of Porocephalus tortus is an interesting subject
for speculation. Possibly both are. It is not so common to find mature Porocephali
in the body-cavity as in the lungs of snakes, but cases of the former habitat are
described. I know no case of the adults being found in the stomach of a host. It
would further seem improbable that parasites subsisting on the healthy, living body
of an animal would desert it to attach themselves to the tissues of another animal
which is not only dead but half-digested. On the whole the evidence points to the
Dipsadomorphus irregularis being the normal host, though the parasites could evidently
live well enough in the gastric juices of Nardoa boa. As a rule Linguatulids endeavour
to quit a dead host and it seems strange that they retained a hold on the swallowed
form so long. I should have expected them to attach themselves to the living host
as soon as the way became open.
Curiously enough although a considerable portion of the anterior end of the
swallowed snake had actually been digested away there were no signs of corrosion at
all on the posterior two-thirds of its body.
The measurements of the fresh specimens of Porocephalus tortus were a little
larger than those I recorded from spirit specimens. Dr Willey gives the length of
the largest as 30 mm., mine was 43 mm. He gives the length of the head as 6 mm.
or including the neck 7 mm., I gave it as 5 mm. He measures the width of the
head at b'b ram. and the width of the body, which he states is " quite round," at
4 mm. I made the latter measurement, taken from somewhat wrinkled specimens,
2'53 mm. Evidently the animals had shrunk in the .spirit.
In my first description of P. tortus I was able to figure three stages, a young
form with some 25 rings and no twisting of the body, an intermediate form in which
the twisting had commenced but the rings were still visible just behind the neck,
and a mature form markedlv twisted ami without trace of rings. I am now able to
give a much better figure of the mature animal taken from a sketch of Dr Willey's
from life, and also to add a couple of figures of two stages younger than the ringed
specimen figured before (Figs. 35, 31 and 32, Plate LYL).
The youngest of these is 2"2 mm. in length, and about 06 mm. in diameter at
the level of its greatest breadth. Behind it tapers to a tail (Fig. 31, Plate LVL).
Some viscera are visible through the semi-transparent skin. In this stage the ringing
DURING HIS SOJOURN IX THE WESTERN PACIFIC. 565
is conspicuous on the tail but comparatively inconspicuous on the trunk. The number
of rings amounted to about forty. These rings are the despair of the researcher on
Lino-uatulids. In the next stage there are at least as many if not more rings
(Fig. 32, Plate LVL), whilst in the third stage (Fig. 83, Plate LVI.) there are but
twenty-five or si.K. I have one specimen killed in corrosive sublimate and not in alcohol
as most of them were, which owing to a fusion of the rings along the median line
has at least twice as many rings on one side of the body as on the other.
The four hooks can just be made out at the anterior end of the second stage,
and in Fig. 32 one is shown in profile, but the}' are much more conspicuous in the
next stage (Fig. 33, Plate LVL).
In the second stage the trunk has encroached on the tail which is beginning
to swell out and consequently to disappear. The rings are beginning to widen out
or possibly two or three of the annulations of the younger forms go to build up
a single one in the older stages. This process has commenced in the stage we are now
dealing with. Its length is o mm. and its greatest breadth is 1'3 mm.
The mature and twisted forms were crowded with the long coiled oviducts twisting
and looping in every direction. The skin was stretched, thin and transparent. The
ova are minute and their number prodigious. With the aid of a haemacytometer I
calculated that there must be at least 3000 to the cubic millimetre, a number which
gives some 825,000 ova in a medium sized adult. There are of course many sources of
error in this method of calculation, but without attaching too great importance to
the figures the calculation serves to show how prolific these animals can be.
List of Ho-sts with their Parasite.s.
Aetiobatis nai-inari, Euphras. Calliobothrium aetiohatis, n. sp. (p. 541).
„ Adelohothrium aetiobatidis, n. gen. (p. 545).
„ „ „ Echinocephalus striatus, Montic. (p. 560).
Histiophorus, sp. Bothyiocephalus pJicatus, Rud. (p. 540).
Pimelepturus fuscxw, Cuv. and Val. Distuinina veiitricosuin, var minor, Pal. (p. 540).
Chelone imhricata, L. Monostomnm trigmocephalum, Rud. (p. 532).
Varanus indicus, Daud. Palaia varani, n. gen. (p. 548).
„ „ „ Fhysahptera varani, Par. (p. 560).
Dipsadomorphus irregularis, Merrem. Phyllobothrium dipsadomorphus, n. sp. (p. 550).
„ „ „ Physaloptera obttisissima, Molin. (p. 559).
„ „ „ „ retusa, Rud. (p. 55.9).
„ „ „ Sclerostomum appendiculatum, Molin. (p. 560).
„ „ „ Porocephalus tortus, Ship. (p. 563).
Carpopliuga van-wycki. Coelodela hmiria, n. gen. (p. 552).
Dioniedea e.iulans, L. Prostliecocotyle diontedeae, n. sp. Fuhrmann (p. 557).
„ „ „ Gnathostoma sliipleyi, n. sp. Stossich (p. 560).
The Zoological Labouatory, December 1899.
Cambridge.
7.5—2
566
A DESCRIPTION OF THE ENTOZOA COLLECTED BY DR WILLEY
EXPLANATION OF FIGURES ON PLATES LIV., LV. AND LVI.
PiEFERENCE LETTERS.
.4. = glandular organ on male ducts.
a.c. = alimentary canal.
£.= dome-shaped organ on male duct.
C.= glandular organ on vagina.
c.s.^. = cavitj' of shell-gland complex.
e.g. = cephahc ganglion.
d.v. = dorsal excretory ves.sel.
e.c. = water- vascular system.
e.ca. =egg cavity.
g.d. = genital ducts.
Z.ra.= longitudinal muscles.
VI, = muscular layer.
M. = longitudinal nerves.
0. = oesophagus.
o.TO. = obUque muscles.
ov. = ovary.
p. = parenchymatous cells.
Pr. glds. = prostatic glands.
»S'. = sucker.
s.g. = shell-gland complex.
t. = testis.
t. r. = transverse excretory vessel.
ut. = uterus.
». = vesicle of water-vascxilar system.
va. = vagina.
v.g. = vaginal glands.
v.d.=va,s deferens.
v.v. — ventral excretory vessel.
W.v. = water- vascular system in head.
y.g. = yolk-glands.
PLATE LIV.
Figs. 1 — 7 inclusive refer to Monostomiim trigorwceplicdum Rud.
Fig. 1. Four sketches showing different aspects of Monostomum trigonocephalum Rud.
1 a. life-size. 1 6. x 8. 1 c. and \ d. x 10.
Fig. 2. Section showing longitudinal and obliquely transverse muscle-fibres. On both sides
of the figure the parenchymatous cells are beginning to appear.
Fig. 3. Longitudinal vertical section tlirough the body slightly to the left of the middle
line.
Fig. i. Section through the glandular structure A, showing the communication of its
lumen with that of the dome-shaped structure B by means of a small muscular valve.
Anteriorly tlie vas deferens is seen cut in many places as it twists and curls in its muscular
sheath.
Fig. .5. A longitudinal horizontal section, showing the loops of the uterus, and parts of
the alimentary canal, the water-vascular system, the testis, shell-gland complex, glandular
organ A, vas deferens and egg-cavity.
Fig. 6. Shell-gland complex. The uterus is shown leaving this organ on the left, and
a coil of it lies to the right. Each contains ova witli deeply stained yolk granules.
Fig. 7. Highly magnified section to show the structure of the peculiar organ C and its
relation to the vagina. The water-vascular canals with their lobes, a piece of the uterus
with the stalked eggs, part of the egg-cavity and some of the vaginal glands are also shown.
Fig. 8. Head of Bothriocephalus plicatus Rud. x about 10. Dorsal or ventral view.
Fig. 9. The same, lateral view.
Results:
Plate LIV
V.dA'.t. W-.^ion Cinbrid^j
.ViLLEY Zoological Results
Plate LV.
SHIPLEY
Edw.n WiUon Cftmbn-igc
DURING HIS SOJOURX IX THE WESTERN PACIFIC. 567
PLATE LV.
Fig. 10. View of the posterior proglottides of B. plicatus. x 8.
Fig. 11. Calliohothrium aetiobatis n. sp. The whole tape- worm, showing end view of
head, x 12.
Fig. 12. Side view of head of C. aetiobatis, showing the bifurcated hooks and tlie three
bothria. x 12.
Fig. 13. Adelobothrium aetiubatidis, n. g. View of the entire worm, x 8.
Fig. 14. End view of head of A. aetiobatidis, showing the rosteUum and the swollen neck
bearing four small suckers on its anterior surface, x 2-5.
Fig. 1.5. A section of the same, showing how the rostellum is completely buried in the
intestinal wall of the Aetiobatin narinari. The swollen neck with two small suckers is shown.
Fig. 16. Head of Falaia varani n. g. surrounded by remains of the lining of the
duodenum of its host Varanus indicus. x 4.
Fig. 17. View of P. vai-ain. x 8. a. Entire worm, showing the grooved nature of the
middle proglottides and some genital pores. 6. Posterior end showing the outline of the
posterior proglottides.
Fig. is. Transverse section of a very young proglottis of P. varajii, showing the deep
longitudinal furrows and the deeply stained longitudinal muscles, and the two longitudinal
water-vascular canals and a commencing genital duct upon each side.
Fig. 19. Transverse section of a ripe proglottis of P. varani, showing the eggs in
clumps in ulero, and the scant remains of the reproductive organs, the penis is partially
protruded.
Fig. 20. A longitudinal horizontal section through ripe proglottides of P. varani, showing
characteristic arrangement of eggs.
Fig. 21. A longitudinal horizontal section near the anterior end of P. varani. x 20.
Showing the characteristic ladder-like arrangement of the deeply staining muscles.
Fig. 22. Head of Phyllobothrium dipsadomorjihi n. sp. x about 12.
Fig. 23. View of head of Coelodela kuvaria n. g. x about 12.
PLATE LVI.
Fig. 23 a. Diagram of the water-vascular plexus in the head of C. kuvaria, seen from
above.
Fig. 24. A longitudinal horizontal section of the adult proglottides of C. kuvaria, showing
the valves in the excretory system.
Fig. 2.0. An obliquely transverse section through older sections of C. kuvaria to show
the great space occupied liy the transverse excretory canal.
Fig. 26. A longitudinal vertical section through C. kuvaria, taken through the most
posterior proglottides.
Fig. 27. Three views of tlie head of Prosthecocotyle diomedeae Fuhrmann. a. Anterior
and dorsal view. h. Lateral view. c. Dorsal view.
Fig. 28. Outline of middle and posterior proglottides of P. diottiedeae.
568 ENTOZOA COLLECTED BY DR WILLEY IX THE WESTERN PACIFIC.
Fig. 29. Section, slightly oblique through the posterior end of the head of P. diomedeae.
The posterior end of the groove of the upper funnel is just visible on the left, the ventral
and dorsal water-vascular systems are shown on each side, those on the left approaching
to fuse with one another. The strongly marked ridges which separate the right and left
suckers are also seen.
Fig. 30. a. View of Qordius rvilleyi Camer. x 1. 6. Head end x about 10. c. Tail
end X about 10. d. The same more highly magnified after Camerano, showing the post-cloacal
lamina and the cloacal opening.
Fig. 31. Youngest known form of Porocephalus tortus Shipley x 10, showing the ringed
tail.
Fig. 32. A side view of a slightly older form of P. tortus x 10, showing an increase
in the number of annulations and a hook ; the head is not constricted from the body.
Fig. 33. A somewhat older form of P. tortus x 2. The head is beginning to separate
off from the body, but as yet there are no hooks. The annulations are very marked, and
the twisting of the body has not begun.
Fig. 34. An older form of P. tortus x 2. The neck has appeared and the growth of
the reproductive organs has stretched the skin so that the annulations have begun to dis-
appear, and the twisting of the body has commenced.
Fig. 35. The oldest stage, side view of P. tortus x 2. The annulations have completely
disappeared, and the twisting of the body is very marked.
Fig. 36. Ventral view of head, neck and anterior end of trunk of the same. From a
sketch by Dr Willey of a live specimen.
WiLLEY Zoological Results.
Plate LVi
Hi'
A E SHIPLEY. ENTOZOA
Edwin Wilson Combridqe
ON SOME SOUTH PACIFIC NEMERTINES COLLECTED BY
DR WILLEY.
By R. C. PUNNETT, B.A.
With Plates LVII.— LXI.
The Nemertines collected by Dr Willey contain representatives of all the four
great classes except the Mesonemertini. Six new species are here described and
mention has been made of at least two other species, which are also probably new.
In addition the collection contains fragments of several others, but as the anterior
end was not preserved in these cases it has appeared wiser to neglect them. A glance
at the map which forms the last plate of Burger's monograph renders it evident
that the Malay archipelago is the headquarters of the genus Eapulia. Consequently
it is not surprising to find that five out of the twelve species mentioned below belong
to this genus. The genus Drepanophorus also has been previously shewn by Burger
to form a conspicuous portion of the Nemertine fauna of this part of the world.
Quite otherwise is the case of Carinesta orientalis which enjoys the distinction of
being the only Protonemertine yet recorded from any place outside the Atlantic and
Mediterranean areas. Cnrinella has been found at Magelhaen's Strait and at the
Cape of Good Hope, but with these two exceptions no member of the class is known
to occur south of the Equator. Its structural peculiarities seem to warrant the
foundation of the new genus; and I may here take the opportunity of thanking
Dr Willey for the chance of examining such an interesting form.
St Andrews, 0<jt. 1899.
PROTONEMER TINP.
Genus. Carinesta, nov. gen.
Head elongated, not spatulate as in the genus Carinella. No cerebral slits present.
The excretory system is simple and is situated some way behind the head. There
is no diagonal muscle layer between the longitudinal and circular layers. No sense
organ near excretory pore.
' The classification followed is that given in Biirger's Monograph.
570 ON SOME SOUTH PACIFIC NEMERTINES COLLECTED BY DR WILLEY.
1. Carinesta orientalis, n. sp. Plate LYII. Figs. 1 — 9 and LVIII. Figs. 10 — 11.
Two specimens were procured from Pigeon Island, New Britain, the larger of
which, though lacking the hind end, measured about 16 cm. in length and was about
2'5 mm. thick. The smaller specimen was still more imperfect, but the whole of the
anterior portion of the body was preserved in each case. The colour was a uniform
dull red-brown, becoming rather lighter towards the anterior end. There were no
markings of any sort.
The mouth is large and is situated shortly behind the brain. The small proboscis
pore is ventral and nearly terminal (Figs. 1 and 11). -The epidermis is not very
thick and contains a number of unicellular glands. Composite glands are absent, and
there are no glands beneath the basement membrane such as occur in the genus
Carinella. Directly beneath the thin basement membrane a very delicate nervous layer
can be made out. In front of the brain this becomes greatly thickened, suggesting
a tactile function for the elongated pre-cerebral portion of the head (Fig. 3, n.l.).
The circular musculature is feebly developed and is lacking in the region of the
brain.
The inner circular musculature which envelops the proboscis sheath and ali-
mentary canal starts behind the movtth. It encloses also the blood vessels at first
but these soon come to lie outside it, and a special portion then surrounds the
proboscis sheath (cf. Figs. 6 and 7). There is a slight dorsal decussation of fibres.
The longitudinal musculature is well developed throughout. The portion surround-
ing the proboscis sheath is throughout the course of the latter well marked off from
the rest, and with the differentiation of circular fibres round it comes to lie external
to these last (Figs. 4, 5, 6 and 7, m.l.jj.s.).
The alimentary canal is straight without any caecum or pouches and appears
relatively large with very high epithelium towards the hinder end of the body.
In the anterior part of the oesophageal region it has special longitudinal muscles
which lie dorsally within and ventrally without the circular muscles (Figs. 5 and 6,
m.l.oes.).
The proboscis is attached just behind the brain (Fig. 11), and possesses a thick
epithelium together with an outer circular and an inner longitudinal muscle layer
(Figs. 10 and 11), the latter being covered by the delicate rh}Tiehocoelom lining.
The two proboscis nerves lie just outside the circular muscles. There are well-marked
glands in the rhynchodaeum near the proboscis pore (Fig. 11, gl-p-).
The vascular system consists of two longittidinal lateral trunks which meet anteriorly
dorsal to the proboscis pore. In front of the brain they give off a ventral and a
dorsal vessel, the former of which joins the loop over the proboscis whilst the latter
ends blindly (Figs. 2 and 3). An anastomosis between the two vessels of the head
loop has been figured by Joubin for Carinella hanyulensis ((6) PI. XXVII. fig. 8),
though it is of a simpler nature and only dorsal. Small vessels are given off to the
oesophagus, but there is no dorsal vessel to the proboscis sheath.
The excretory system is small and consists of a glandular portion lying inside
the lateral blood vessel as in Carinella, and a small duct opening more posteriorly
ON SOME SOUTH PACIFIC XEMERTIXES COLLECTED BY UK WILLEY. 571
to the exterior (Figs. 2, 7, and 9). There is no direct communication between
any part of the vascular and excretory sj'Stems. The excretory system is relatively
further behind the brain than in any other Nemertine.
The generative organs were not quite ripe. They formed a continuous band on
each side of the body. Doubtless thej^ represented a number of glands closely apposed
and each opening by its peculiar orifice. These however were not yet distinctly formed,
though traces of them could be made out here and there (Fig. S, ov.).
The brain is simple in structure and has the appearance of the nerve cords being
much swollen in this region, since the dorsal ganglion is not sharply separated off
from the ventral. There are two dorsal ;in(l two ventral commissures, the posterior
dorsal and the anterior ventral lying at the same level (Fig.s. 2 and 4). From
the posterior ventral commissui'e are given off the two strong oesophageal nerves.
There is a small median dorsal nerve extending backwards just beneath the
basement membrane. In front of the brain there is a well-marked nervous layer
just beneath the basement membrane (Figs. 3 and 11, n.l.). There are no neuro-
chords.
There are no special sense organs, eyes and frontal organ being absent. More-
over there is no lateral sense organ such as has been described by Burger for
Carinella, in the neighbourhood of the excretory pore.
Carinesta orientalis is one of the few Nemertines without cerebral slits, a peculiarity
shared with CarinovHO and Malacohdella alone.
jVETA nemer tixi.
'1. Drepiinophorus willeyanus, n. sp. PL LVIII. Figs. 12 — 17 antl LIX. 18 — 20.
A single specimen was found by Dr Willey in Blanche Bay. Its dimensions when
preserved were 7'5 cm. in length and 2'3 mm. in breadth. The posterior end was
sharply pointed. In life the worm was of a lightish brown dorsal ly whilst the
ventral surface was dirty white. The head furrows were very conspicuous in
the living animal as a series of deep chocolate brown stripes on either side dorso-
laterally (Fig. 12). They are more strongly developed than in any other .species of
Drepanophonis, their condition being most nearly approached by that obtaining in
B. spectabilis. The eyes were easily seen in the living animal and were disposed in
the arrangement characteristic of the genus. Uorsally and laterally there was a row
of three eyes on each side whilst ventro-laterally and rather more anteriorly was a
row of five on each side (Figs. 12 and 13), thus making a total of sixteen eyes.
All the other known species of Drepanophorus possess at least thirty eyes, and this
character has been taken by Burger as characteristic of the genus (Biirger (4)
p. I(i8), so that in this respect D. willeyanus differs markedly ho\n its relatives.
Tlie basement membrane, as is usual in this genus, is well marked, being nearly
as thick as the epidermis (Figs. 13 and 18, b.m.). The body parenchyma in which
' Biirger states that there are no cerebral slits in Cephalothrix and makes this a character of the genus.
Joubin however figures them for Cephalothrix linearis ((8) PI. XXVI. tig. 24).
w. V. 76
572 ox SOME SOUTH PACIFIC NEMERTINES COLLECTED BY DR WILLEY.
the various organs lie is well developed, aud is traversed by uumerous dorso-ventral
muscle strands which are regularly arranged laterally and separate the various organs
which are subject to repetition (Figs. 15 and 16, in.d.v.).
The muscles of the body wall present the characteristic arrangement of the genus,
and are not very strongly developed.
The oesophagus is small. Shortly after the brain region and before the termination
of the cerebral organ it dilates into the mid-gut (" Magendarm ' ). The blind gut
commences anteriorly at the level of the second rhynchocoelom diverticulum aud extends
backwards to the genital region, where it joins the main gut at the junction of the
now somewhat attenuated raid-gut and intestine proper. The blind gut gives off lateral
pouches and in histological features resembles the intestine which also gives off
pouches along its course. The anus is terminal though slightly ventral in position.
The proboscis is stout and is covered with papillae which are supplied by strong
branches from the proboscis nerves (Fig. 18). Beneath the papillated epithelium is a
layer of circular muscles and then a strong layer of longitudinal muscles which is
divided into two portions by the ring formed by the proboscis nerves and their
commissures. The proboscis nerves are 16 in number, which is probably a characteristic
of this species.
The musculature of the proboscis sheath shews the basket-like structure characteristic
of the genus (cf. Hubrecht (6) p. 106 and PI. X. figs. 4 and 5, and Burger (1)
Fig. 1.50).
The rhynchocoelom diverticula are very large and spacious extending round the
gut-pockets ventrally and with great regularit}-. The size of their communications with
the rhynchocoelom differs greatly, being in some cases very wide, especially towards
the anterior end, whilst in others they can hardly be distinguished. There is no
connection between neighbouring diverticula although the}" may be in contact anteriorly
in the pregonidial region.
The vascular system is of the ordinary type described by Oudemans for Drepano-
phorus (Oudemans (9) PL I. fig. 7).
The nervous system resembles closely that described by Biirger for D. spectabilis.
The dorsal and ventral commissures are of approximately the same size. A number
of strong nerves are given off anteriorly to supply the large eyes. The lateral nerv^es
-give off commissures which pass ventral to the gut. They also give off branches
passing dorsally, but these do not effect a junction with those of the opposite
side such as Hubrecht has described in the case of Drepanophorus lankesteri (Hubrecht
(6) p. 107).
A fact well wnrthy of note is that the repetition of the ventral commissures in
this species does not correspond with the repetition of the intestinal pouches, gonads,
and muscle septa, in the way that Blh-ger has- figured for D. cnissus (Burger (:3)
Taf 17, fig. 16).
The cerebral organ is large, being almost as bulky as the cerebral lobes. It
commences just behind the brain with which it is connected by a single strong nerve
and not by three as in D. spectabilis. The posterior glandular portion is of the same
length as the organ itself, being intermediate in this respect between B. spectahilis and
D. cerinus. The oesophageal nerves are small.
ON SOME SOUTH PACIFIC NEMERTINES COLLECTED BY DE, WILLEY. 573
The excretory system reaches anteriorly almost as far as the hind end of the brain.
Posteriorly it stretches nearly as far as the commencement of the gonads. There is a
single pore on each side which occupies a very forward position at the level of the
conunencement of the mid-gut and in front of the hinder end of the cerebral organ.
The generative organs were nearly ripe. The individual is a ?. The ovaries are
large and crescentic in shape, and each with a single opening (Fig. 20). They alternate
regularly with the intestinal pouches and rhynchocoelom diverticula from which they are
separated by muscle strands.
Thus it will be seen that this species of Drepanophorus is well marked from
other species of the genus, being characterised by the unusually large head furrows,
the small number of the eyes, the large size of the rhynchocoelom diverticula, and by
the number of the proboscis nerves. I have much pleasure in associating it with
Dr Willey's name, more especially since he examined the animal carefully when alive
and made the interesting observation that "when just taken it frecpiently and rapidly
alternately extruded and retracted its probo.scis much in the same way that Ctenoplana
uses its tentacles."
3. Drepanophorus lifuensis, n. sp. PI. LIX. Figs. 21 — 27.
The single specimen of this species which was procured at Lifu is a flattened form
measuring 4 mm. in width, 1'5 mm. in depth, and 4'5 cm. in length. A coloured sketch
was made b}' Dr Willey of the live animal from which it appears that the dorsal
surface was of a bright palish red, whilst the ventral was quite white. The anterior
end was whitish and on it the eyes appeared as four dark blue longitudinal lines. The
appearance of the preserved animal agrees closely with that given by Burger for
D. Iritus ((1) p. 27). The animal possessed a small " tail," coloured like the rest of the
body (Fig. 26, c). Sections made through it shewed that the anus opened at its
extremity. There was, however, no anal commissure, the nerve cords ending blindly
and somewhat abruptly, and I am inclined to believe that this " tail " has its origin
in some regeneration process following upon an injury to the posterior end of the
w'orm.
Another point of interest with regard to the external features of the animal lay
in the fact that the epidermis, with the exception of a few small isolated patches
here and there, had almost entirely disappeared, the basement membrane being left
exposed to the external woi-ld. Coupled with this was the presence of a small
insect larva a few millimetres in length, labelled by Dr Willey as parasitic on this
worm. One is tempted to draw the conclusion that this little creature obtained its
nourishment by browsing upon the epidermis of the Nemertine, in which case it would,
with the exception of latoshia gigas, be the only external parasite recorded for the
group.
With regard to internal features the species shews a great resemblance to D.
lutus.
The head glands (" Kopfdriisen ") are short and feebly developed. The mouth is
slit-like and ventral in position, and the proboscis pore is at its anterior end. The
70—2
574 ON SOME SOUTH PACIFIC NEMERTINES COLLECTED BY DR WILLEY.
oesophagus passes into the mid-gut in the posterior brain region (Fig. 23), though before
this happens a well-marked typhlosole-like process is found projecting from it in the
mid-ventral line (Fig. 23, oes.t.).
The proboscis shews the structure typical for the genus and contains 31 nerves.
The excretory pore is situated "35 mm. behind the cerebral organ. The whole
extent of the excretory system is about 2'2 mm. of which one-seventh lies in front of
the pore, so that the excretory tubules do not reach quite to the cerebral organ. Near
the pore the excretory duct dilates into a bladder.
The diverticula of the proboscis sheath commence at the termination of the cerebral
organ. They are rather narrow and reach round the intestinal pockets about half-way
between the nerve cords and the mid-ventral line.
The brain is large and in shape differs greatly from that described by Burger
for D. liitus. Instead of being " schlank, elliptisch, von geringeren Querdurchmesser,
aber aussordentlich hoch," it is well rounded (Fig. 23). The ventral commissure is
extraordinarih' strong (Fig. 22).
The cerebral canal opens exactly laterally, not somewhat dorsally as in D. latits.
The cerebral organ is large and ends at the same level as the dorsal ganglion, i.e.
no part of it lies behind the brain. It possesses a layer of gland cells on its ventral
surface, which glandular structure does not project backwards from the rest of the
organ as in some other species, e.ff. D. willeyanus (Fig. 19, gl.). The pigment masses
in the cerebral organ are comparatively very large (Figs. 24 and 27), and were visible
in the living animal (Fig. 26, c.org.).
The parenchyma is well developed, being particularly dense where it surrounds
various organs such as the nerve coi'ds, excretory tubules, etc. (which indeed have
shrunk away from it in conservation). In the light of Dr Willey's observation that
this animal swims about freely such a fact becomes of interest when we regard such
a form as Pelagonemertes where free-swimming pelagic habits are accompanied b}'
an enormous development of the parenchyma. Possibly the substance serves to take
the place of a skeletal structure enabling the relatively slightly developed muscular
layers to bring about sharjj vigorous contractions in definite directions resulting in
forward propulsion.
HETERONEMERTINI.
4. Eupolia rugusa, n. sp. PI. LIX. Fig. 2S and LX. Figs. 29—31.
A single specimen was taken at Xew Britain. It measured about 3 cm. in
length (Fig. 31) and the skin in the preserved specimen was deeply wrinkled. Dr
Willey tells me that these wrinkles were present when the worm was living, and I
have accordingly given it the name which stands at the commencement of this
description. In the characters and ari-angement of the skin, cutis, and muscle layers
the animal presents no marked divergence from the general type of the genus.
The head glands are enormously developed. They open at the anterior margin
of the proboscis pore which is ventrally situated. Lying in the outer longitudinal
ON SOME SOUTH PACrFIC NEMERTINES COLLECTED BY DR WILLEY. 575
muscle layer they stretch tar past the brain into the mid-gut region. Some idea of
their extent may be gathered when it is stated that in the series of sections through
the animal the brain occupies sections 90 — 135 whilst the head glands reach to
section 2G0. They are rendered exceedingly conspicuous by thidiiin in which they
take a deep reddish purple tinge (Figs. 2.S and 29).
The head slits are deep and ventral in position reaching from the anterior end
to a short distance behind the spot where the cerebi'al canal opens into them
(Fig. 28).
The dorsal ganglion passes insensibly into the cerebral organ, the glandular
layer of which lies quite internal (Fig. 30). Inside this glandular portion were a
number of small circular bright green bodies containing in the centre a well stained
nucleus. Their brilliant colour renders them exceedingly conspicuous but I am unable
to make any suggestion as to their nature. In appearance they bear some resemblance
to unicellular algae. Moreover many of them are clustered together suggesting that
some process of division had been going on (Fig. 30 *j. The nerve cords form no
commissure by the anus.
The circular head furrow so characteristic of the genns appears to be absent.
The excretory system is continuous and opens by several ducts on each side,
though these are fewer in number than in E. multiporata. The ducts pass as usual
just over the dorsal side of the lateral nerve cord.s.
About the vascular system nothing definite can be said as the state of con-
traction rendered it impossible to observe with any certainty the course of the vessels
from section to section. No eyes are present.
5. Eupolia heinprichi (Ehrenbekg, 1831). PI. LX. Fig. 32.
A single specimen from sand in rock pools at Lifu. With regard to its markings
the animal shews a slight divergence from that described by Burger (1) page 22
(under the name of E. brockii) in that the brown transverse line near the anterior
extremity is not continued ventrally to form a complete ring. Also the ventral brown
line commences further back and soon becomes indistinct. The length of the animal
in life was over 30 cm., its breadth being about 3 mm. of which 1 mm. was taken
up by the dorsal longitudinal stripe. A figure of the animal is given in the paper
cited above, Taf I. fig. 10.
With regard to its inner organisation it agrees with Btirger's account. Although
I have a complete series of sections through the anterior portion of the animal
I have been unable to find the openings of the excretory system which in this genus
are generally so well marked as they traverse the thick gelatinous layer of the cutis.
An interesting point is the presence of large cells resembling ova scattered about
and lying as far forwards as the level of the cerebral organ (Fig. 32, ov.). As each
is surrounded b}' a fine epithelium with fiattened nuclei, and lies in a cavity lined
by a similar epithelium, it seems more natural to regard them as ova than as parasitic
Gregarines. Unfortunately the animal contained no undoubted ova with which to
compare them.
576 ON SOME SOUTH PACIFIC NEMERTINES COLLECTED BY DR WILLEY.
6. Eupolia quinquelineata (Burger, 1893). PI. LX. Fig. 33.
Two specimens were obtained from different localities.
The first which was found at New Caledonia measured after preservation
in length .52 cm.,
„ breadth 12 mm.,
„ depth 6"o mm.
In the arrangement of the dorsal stripes this specimen differs somewhat from
that figured by Biirger (2). The two outer lines were very imperfect and over con-
siderable portions of the body were entirely absent. ' The two inner lines were also
interrupted at intervals. The median dorsal line was broken in one place near the
head. Willey describes the dorsal intervening ai'eas as " opaque white with faint
crimson tinge locally." The two ventral stripes were not interrupted, and were each
located " in the middle of a dull slightly pinkish subtranslucid belt." About the
middle of the body an injury had evidently occurred and been repaired. Fig. 33
shews the effect on the arrangement of the lines.
The second specimen, which came from New Britain, agrees in the arrangement of
the stripes with that figured by Biirger. Its dimensions when preserved were
length 45 cm.,
breadth 10 mm.,
depth o mm.
The posterior end was however broken.
t.
Eupolia midtiporata, n. sp. PI LX. Figs. 34—39 and LXI. Fig. 40.
A single specimen from Rakaiya, New Britain. The animal was rounded in shape,
the length being 12 cm. and the breadth 3'8 mm. The preserved animal is of a pale
green colour and shews no markings. The mouth and proboscis pore shew clearly
(Fig. 34) ; the circular head furrow is complete and well marked ; the posterior
extremity is sharply pointed.
The eyes are small and numerous and are all situated in front of the head
furrow (Fig. 35, oc).
The skin is of the usual Eupolia type with the slight differences that the deep
glands of the cutis have very broad bases and that the epidermis contains small
compound glands (Fig. 39, gl.d and gl.c).
The head glands are practically absent', a point in which this species shews a
marked difference from the rest of the genus. The outer longitudinal muscle layer
extends to the tip of the suout. Here it is much broken up (Fig. 36). The circular
and inner longitudinal layers do not appear until the level of the cerebral organ and
then only on the dorsal side. The relative thickness of the different muscular layers
(and also of the cutis^) is depicted on Fig. 40.
1 Biirger (3, p. 179) writes "Die Eupolien sind alle ausgezeichuet durch eine machtig entwickelte Kopfdriise.
ON SOME SOUTH PACIFIC NEMERTINES COLLECTED BY DR WILLEY. 577
The proboscis sheath is small and of the usual type. Just over the proboscis
vessel the epithelium becomes high and glandular (Fig. -iO, psjjl.). 'J'he proboscis
is lost
The wall of the mid-gut consists of two well-marked portions, the inner ciliated
epithelium and the outer glandular portion, such as has already been described by
Hubrecht for one of the "Challenger" species {E. nipponensis). The vascular system
consists in the snout of wide laterally extending lacunae lying dorsal to the proboscis
sheath (which here of course is morphologically proboscis epithelium of ectodermal
origin) (Fig. 36, b.v.). In the brain region the vascular system comes to lie ventral
to the proboscis sheath and comprises a large median and two small lateral vessels
(Fig. 37). Further back the lateral vessels become enlarged and sink down towards
the oesophagus (Fig. 38), in which region they break up into the plexus so characteristic
of the genus (Fig. 40).
The excretory system is peculiar. Not only are there a great number of ducts
(probably 20 — 30 on each side), but the excretory gland is broken up into a immber
of portions. Numerous ducts have been described for E. (jiardii by Hubrecht (6) and
for E. curta by Oudemans (9), but the gland was always continuous. The only other
case of a discontinuous gland among the Heteronemertini is that recorded by Coe (5),
p. 498, for Valencinia, but the phenomenon was far less marked in that case. A
reconstruction of the system is given in Fig. 35. Unfortunately this is not quite
complete as some sections containing the most posterior portion were lost. It could
not have extended much further backwards however as it is not found in sections
about 2 mm. posterior to the last shewn on the reconstruction. Hubrecht ((6), p. 116)
lays some stress on the correspondence of the ducts on the two sides. It will be seen
that practically none exists here.
The brain is small for the size of the animal, and the dorsal ganglion is larger than
the ventral.
The ventral commissure is strong (Fig. 37).
The cerebral organ contains a pigment mass. It communicates with the exterior in
the head furrow ventrally (Fig. 37). The peculiar epithelium of the ciliated canal of the
cerebral organ is continued into and all round the head furrow (Fig. 37, ep.h.f.).
8. Eupolia, sp.
Fragments of a large white Eupolia were found by Dr Willcy living in sand
together with Ptychodera at Rakaiya. As the anterior portions of both the specimens
preserved were unfortunately missing it is impossible to give any further account of
the species.
9. Linens au.stndis, n. sp. PI. LXI. Figs. 41 — 44.
A single specimen from Lifu. The jtosterior portion of the body is broken off, so
that it is impossible to say whether it possessed the small tail which forms the chief
criterion, as regards external features, between the genera Lineits and Cerebratulus.
The structure of the skin however, which resembled that of the genus Eupolia,
probably settles its position in the genus Lineus.
578 ON SOME SOUTH PACIFIC NEMERTINES COLLECTED BY DR WILLEY.
The colour of the preserved animal was a uniform dirty brown. A lighter area
extended along the edges of the head slits and round the proboscis pore (Fig. 41).
The mouth commenced after the posterior end of the head slits. No eyes are present.
The head glands are fairly strongly developed, extending for two-thirds of the
distance between the brain and the tip of the head, where they open by several
small pores.
The skin greatly resembles that of the Eupolidae in structure, the gelatinous layer
of the cutis (" Bindegewebeschicht " of Burger) being, at the end of the raid-gut region,
as thick as the rest of the cutis and epidermis together (Fig. 42). A similar condition
has been described by Blirger for L. geiiiculatus.
With regard to the vascular system the two loops of the head-ring unite just in
front of the brain (Fig. 43 (a)). Just anterior to the level where the cerebral organ is
first seen and where the lateral nerves are given off the single vessel so formed divides
into four (Fig. 43 (c)), a ventral and a doi-.sal median vessel and two lateral vessels.
These two lateral vessels then take an upward course over the dorsal ganglion and,
after surrounding it, unite ventrally with one another and with the median ventral
vessel (Fig. 43 (e)). The median ventral vessel here disappears and the lateral vessels
form large sinuses reaching more than half-way round the cerebral organ (Fig. 43 (f)).
From these sinuses the lateral blood vessels of the trunk are eventually given off.
Meanwhile the median dorsal vessel enters the proboscis sheath (Fig. 43 (f)).
The excretory pore opens very shortly after the termination of the cerebral organ,
opening laterally from an excretory duct which passes just over the lateral nerve.
Only one pore occurs on each side. The excretory tubules all lie anterior to it and
on the dorsal' side of the great blood sinus surrounding the cerebral organ. The
whole extent of the system is small being only '3 — "4 mm. in the contracted
specimen.
The head slits are very deep, the distance between their termination and the
brain being only one-twelfth of the depth of the slit. The general characters of the
brain call for no special comment. They are shewn in (Fig. 43 (a) — (f)).
A frontal organ is not present.
10. Li'neus alho-vittutus BuRGER, 1890. PL LXI. Figs. 46—47.
A single specimen lacking the hind end was taken at Lifu. Length about 18 cm.
when alive. The colour of the living animal was a dark black green. On the dorsal
surface of the head there was a zigzag line of a pale greenish white hue (Fig. 47).
In spirit the appearance of the animal greatly resembled the figure given by Burger
((1) PI. I. Fig. 1), being characterised by the small size of the head and the enormous
mouth, as well as by the colour and head marking.
In internal anatomy it also agrees with Biirger's description, the peculiar small
' A dorsal position for the excretory system has been described by Biirger for L. gilvus, L. pdrviilus, and
L. gesserensis, but in all these cases the excretory pore is not lateral but dorsal in position. Moreover the
species here described differs from all tliese three in various points, e.g. structure of skin, size of head slits,
size, absence of eyes, &c.
ON SOME SOUTH PACIFIC NEMERTINES COLLECTED BY DR WILLEY. 579
ganglia which lie outside and above the dorsal commissure and supply the head slits,
being also well marked in this specimen. I have, however, been unable to discover the
eyes mentioned by Burger. The following points may now be added to the previous
description.
The head glands are numerous, but fine and scattered. They do not reach to the
brain. The excretory system extends to the posterior limit of the mouth region. The
tubules reach up from the level of the nerve cords to the proboscis sheath. They com-
municate with the exterior by a number of large ducts on each side. The blood
vessels in this region are exceedingly minute. The head slits reach almost to the brain.
The species occurred in holes in coral stone and fixed itself in cavities of the stone
by rendering itself extremely varicose (Fig. 46).
11. Lineus, sp. PI. LXI. Fig. 45.
This species has only reached me in the form of a coloured sketch. It was
found coiled up inside coral stone at Lifu. In length it measured about 8 cm. The
colour was dark green, the tip of the head being white. There was also a white
w-shaped line on the head. The green of the body was relieved by 41 white trans-
verse stripes. Several eyes occur on the outer sides of the green patch at the tip
of the head (Fig. 45). In general appearance this worm greatly resembles L. geni-
culatus, but differs from it, (1) in its smaller size, (2) in the greater number of the
white bands, (3) in the absence of any red colouration round the head slits, (4) in the
eyes not being placed on the w-shaped white band.
12. Langia, sp. PI. LXI. Fig. 4S.
The posterior end of a fragment belonging to this genus was preserved. This
fragment measured about 8 cm. in length and 5 mm. across at its broadest part. The
small tail was present. The whole of the worm in front of the intestinal region was
lost. The colour was a pale reddish brown, the tail being quite white. The worm may
possibly be identical with the L. ohockinna of Joubin (7), in spite of the fact that his
species had no tail. That may very easily have been broken off". Fig. 48 shews a
transverse section of the worm. It was found in Gavutu, New Florida.
w. V.
580 ON SOME SOUTH PACIFIC NEMERTINES COLLECTED BY DR WILLEY.
PAPERS REFERRED TO.
1. Burger, O. Untersuchungen iiber Anatomie und Histologic der Xemertinen. Zeit. fiir
wiss. Zool., 1890.
2. Burger, O. Sudgeorgische und andere exotische Nemertinen. Zool. Jahr. Abt. fiir Syst.,
1893.
3. Burger, O. Die Nemertinen. Fauna und Flora des Golfes von Neapel. Berlin, 1895.
4. Burger, O. Die Schnurwiirme. Bronn's Tierreich.
5. Coe, W. R. On the Anatomy of a species of Nemertean, A-c. Trans. Connect. Acad.,
1895.
6. HuBRECHT, A. A. W. Report on the Nemertea collected by H.M.S. Challenger. London,
1887.
7. JouBiN, L. Note sur 1 Anatomie d'une Nemerte d'Obock. Archiv. Zool. Exp., 1887.
8. JouBiN, L. Recherches sur les TurbeUari^s des c6tes de France. Arcliiv. Zool. Exp., 1890.
9. OuDEM.\NS, A. C. The circulatory and nephridial apparatus of tlie Nemertea. Quart. Journ.
Mio. See, 1885.
WlLLEY Z OOLOGICAL ReSULTS.
Plate LVir
n..l.p3.
<L. b ■
:,iLx
,.ep
\-PP
,<^
oL^ post
% V
''% ..// /
-frv
/ — n.c
Wcet^wraan hth
PUNNETT NEMERTINES.
ON SOME SOUTH PACIFIC NEMERTINES COLLECTED BY DR WILLEY.
581
EXPLANATION OF LETTERS USED IN PLATES.
a.c anal commissure
b.m basement membrane
b.v lateral blood vessel
c.c cerebral canal
cer cerebrum
c.org cerebral organ
cv cutis
cu.gel gelatinous layer of cutis
d.b.v dorsal blood vessel
d.c dorsal commissure
d.c.ant anterior dorsal commissure
d.c. post posterior dorsal commissure
d.g dorsal ganglion
d. n dorsal nerve
d.v dorsal blood vessel
ep epidermis
ep.h.f. epithelium of head furrow
ex.bl excretory ' ' bladder ' '
ex.d excretory duct
ex.gl excretory gland
ex.p excretory pore
ex. t excretory tubules
g.c ganglion cells
gl glandular portion of cerebral organ
gl.c composite epidermal glands
gl.d deeper cutis glands
gl.p proboscis pore glands
h.f. head furrow
h.g head glands
h.s head slit
int intestine
int. d intestinal diverticulum
m mouth
m.c circular muscle layer
m.ci internal circular muscle layer
m.d.v dorso-ventral muscles
m.g mid-gut
mg.gl glandular tissue of mid-gut
m.l longitudinal muscle layer
m. I. ext external longitudinal muscle layer
m. I. int internal longitudinal muscle layer
m. I. oes longitudinal muscle layer of oesophagus
m.l.p.s longitudinal muscle of proboscis sheath
n. c lateral nerve cord
n.cl nerve cells
n.c.org nerve to cerebral organ
n.l nervous sub-epidermal layer
n.oc nerves to eyes
n.oes oesophageal nerve
oc eye
oc.d dorsal eye
oc.v ventral eye
oes.t oesophageal "typhlosole"
ov ovary
ov.d dorsal portion of ovary
ov.v ventral portion of ovary
p proboscis
par parenchym
p.ep proboscis epithelium
pg.m pigmented mass of cerebral organ
p.n proboscis nerve
p.p '....proboscis pore
p.s proboscis sheath
ps.gl glandular thickening of proboscis sheath
TCt proctodaeum
rh rhynchocoelom
rh.d rhynchocoelom diverticulum
r.gl rhynchodaeal glands
V. b.v ventral blood vessel
v.c ventral commissure
v.c.ant anterior ventral commissure
v.c. post posterior ventral commissure
PLATE LVII.
Fig. 1. Carhiesta orientalis. Sketch of anterior extremity, after preservation, x 6.
Fig. 2. C. orieyitalis. Schematic reconstruction, from a series of sections, shewing the
arrangement and relations of the nervous, vascular, and excretory systems.
Fig. 3. C. orientalis. Section through precerebral region, x 110.
Fig. 4. C orientalis. Section through brain, x 110.
Fig. 5. C. orientalis. Section through mouth region, x 60.
Fig. 6. C. orientalis. Schematic representation of the arrangement of the various muscle
layers shortly behind tlie mouth.
77—2
582 ON SOME SOUTH PACIFIC NEMERTINES COLLECTED BY DR WILLEY.
Fig. 7. C. orientalis. Section in the neighbourhood of the excretory pore, x 45.
Fig. 8. C. orientalis. Section through posterior region of body, x 60.
Fig. 9. C. orientalis. 9 a shews the communication of the glandular portion of the ex-
cretory system with the duct. 9 6 is taken a few sections posteriorly to 9 a.
b.io. wall of blood vessel.
exc.d. excretory duct.
exc.gl. glandular part of excretory system.
PLATE LVIII.
Fig. 10. C. orientalis. Section through unextruded proboscis, x 110.
Fig. 11. C. orientalis. Longitudinal section through anterior end. Reconstructed from
several adjacent sections, x 60.
Fig. 12. Drepanophm-us tvilleyanus. Sketch shewing anterior end of animal. Slightly
modified from a sketch made during life by Dr Willey. The only colour is chocolate brown.
X about 12.
Fig. 13. Z>. willeyanus. Section through anterior end shewing the position of the eyes, x 60.
Fig. 14. D. icilleyanus. Section through brain and head furrows, x 45.
Fig. 15. D. ivilleyanus. Longitudinal vertical section through the body near the lateral
edge. Cf. Fig. 18 a... a.
Fir,. 16. D. mlleyanus. Section taken in same plane as last but rather nearer the middle
line. Cf. Fig. 18 /3...y8.
Fig. 17. D. willeyanus. Section shewing supra-anal commissure, x 110.
PLATE LIX.
Fig. 18. Z>. iviUeyanus. Section through middle of body, x 45. * marks opening of
rhynchocoelom diverticulum.
Fig. 19. £>. iviUeyanus. Reconstruction shewing relations and extent of cerebral organ
and excretory system. x 40. * level where rhynchocoelomic diverticula commence. t level
where gut pouches start.
Fig. 20. Z>. willeyanus. Section shewing shape and opening of an ovary, x 45.
Fig. 21. Drepanophorus lifuensis. Section through precerebral region just behind common
opening representing proboscis pore and mouth, x 60.
Fig. 22. D. lijuensis. Section through brain shewing the extremely large ventral com-
missure. X 60.
Fig. 23. D. lifuensis. Section through cerebral region just anterior to where the lateral
cords leave the brain, x 60.
Fig. 24. D. liftiensis. Section through region of cerebral organ, x 60.
Fig. 25. D. lifuensis. Section through level of excretory pore shewing the dilated "bladder."
X 45.
OlOAL RE:
Plate LVIII,
14
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bm^
I k
\-^.
SaSEMv^^"
^- VMHi- ■/
post
/
mcps
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PUNISTETT T-JEMERTINES.
W-a':,Nev/nian iltc
Wn.LEY
OICAL Bes
Hy
\r
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j:t. nci
p 'T ■-- ■
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ON SOME SOUTH PACIFIC NEMERTINES COLLECTED BY DR WILLEY. 583
Fig. 26. D. lifuensis. Shewing anterior end (dorsal surface a and ventral surface 6) after
sketches made by Dr Willey from life. The longitudinal lines at the anterior extremity were
deep blue, the rest of the shaded portion pale red, the unshaded portion white. 26 c shews the
" tail."
Fig. 27. D. lifuensis. Sketch of anterior end as seen when cleared in xylol.
Fig. 28. Eupolia rugosa. Section through brain region. Owing partly to the great state
of contraction, and partly to the strong ventral curvature following tixation, the proboscis sheath
is cut three times and a portion of the mid-gut has come to occupy a position just dorsal to
tlie brain. For the same reasons a portion of the ganglion cells of the lateral cord is also
seen in the section, x 45.
PLATE LX.
Fig. 29. E. rugosa. Section through anterior edge of proboscis pore shewing the openings
of the largely developed head glands, x 45.
Fig. 30. E. rugosa. Section through cerebral organ. Beneath it is seen the lateral nerve
cord passing outwards. * denotes the small green bodies lying in the glandular part of the
organ (vide Text, p. 575).
Fig. 31. E. rugosa. Shewing animal preserved. Natural size.
Fig. 32. Eupolia hemprichi. Section through cerebral organ shewing ovum lying dorsal
to it on one side.
Fig. 33. Eupolia qainquelineata. (a) Dorsal surface of anterior end ; (b) ventral surface,
the shaded portion round the black lines being of a pinkish hue in life ; (o) shews arrangement
of lines dorsally at point of injury (?) about half-way down the animal ; (d) is ventral surface
of the same. All x | (about.) From sketches made by Dr Willey during life.
Fig. 34. Eupolia multiporata. Anterior end. x 4.
Fig. 35. E. midtiporata. Reconstruction shewing position and arrangement of excretory
system. The duct marked * was double, one part opening more dorsally than the other.
Fig. 36. E. multiporata. Section through portion of head anterior to head furrow, x 60.
Fig. 37. E. multiporata. Section through cerebral region, x 60.
Fig. 38. E. multiporata. Section through posterior brain region, x 60.
Fig. 39. E. multiporata. Section through skin in mid-gut region shewing the broad bases
of the deeper glands, x 160.
PLATE LXI.
Fig. 40. E. multiporata. Section through mid-gut region. One of the excretory ducts is
seen passing to the exterior, x 60.
Fig. 41. Lineus australis. Sketch of dorsal («) and ventral (6) surfaces of head end as
seen after clearing in xylol, x about 10.
584 ON SOME SOUTH PACIFIC NEJIERTINES COLLECTED BY DR WILLEY.
Fig. 42. L. australis. Section through commencement of intestinal region shewing the
relative thickness of the various elements of the body wall, x 60.
Fig. 43. L. australis. a^". Sections through brain shewing arrangement of vascular
system in this region. * in / indicates the position the excretory tubules occupy a few sections
more posteriorly.
Fig. 44. Z. australis. Section through skin shelving thickness of gelatinous layer of the
cutis. X 160.
Fig. 45. Lineus sp. 2 After a sketch made of the living animal by Dr Willey. The
ground colour is dark green, and is broken by 41 transverse bands, and by an oj-shaped line
on the head. Natural size.
Fig. 46. Lineus albovittatus. Shewing the varicosity produced at the posterior end by
which the animal fixes itself in holes in coral rock. After a sketch from life by Dr Willey.
Much reduced.
Fig. 47. L. albovittatus. Enlarged -^-iew of anterior end of animal when alive ; after
sketch by Dr Willey.
Fig. 48. Langia sp. ? Transverse section, x 20.
WiLL.BT Zoological Results,
Plate LX.
33
ib)
!
Ill
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If
i
30
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35
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36
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34
PUNNETT NEMERTINES.
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(^
ON THE YOUNG OF THE ROBBER CRAB'.
By L. a. BORRADAILE, M.A.,
Lecturer of Selwyn College, Cambridge.
In October 1874, when the 'Challenger' arrived at Zamboanga in the Philippines,
von Willemoes Suhm was anxious to investigate the development of Birgus latro from
the egg. Unfortunately for his intentions the breeding season of the animal is, in
that part of the world, the month of May. He made enquiries, however, on the subject,
and was told by an " intelligent native " that the young are born resembling the
parent. The robber crab being a land animal coming of a sea stock, this was not
a priori unlikely, though the small size of the female opening would not allow of a
very yolky egg. The suggestion has accordingly been accepted as likely by sundry
text-books.
No further information on the subject has been forthcoming till the recent
publication by the present writer of a short note, showing that the young are hatched
in the zoaea stage (20).
The facts of the case are as follows :
The breeding season in the Philippines is the month of May, and in the Loyalty
Islands January and February. In Christmas Island it is also at the beginning of
the year. At this time the crabs, which ordinarily live more inland'^ may, in Lifu, be
' Birgus latro (Linn.), 1767. (2, 15.)
- The frequency of the visits of Birgus to the sea is still a matter of doubt. According to Darwin (8)
the crab is .said to go down to the shore every night "no doubt for the purpose of moistening its bratichiuf."
Von Suhm was told at Zamboanga that the creature "occasionally" visits the sea (17). Guppy (11) and
Whetham (10) also mention this habit. On the other hand Andrews (22) knows nothing of it, and Dr Willey
tells me that, out of the breeding season, Birgus is taken only inland and at night by torchlight, or can
be trapped in the same place by a cage baited with roasted coconut. The holes in whicli it lives are, at
Lifu, inland under coral limestone. At Zamboanga they are said to be .situated at the roots of trees in
swampy ground. It seems pretty certain that the principal food of the animal is the fruit of the Pandanus
and Coconut trees (6, 8, 14, 19, etc.) which it can obtain, if necessary, by climbing for it, and in this case
there can be little doubt that its object in visiting the sea out of the breeding season would be to moisten
its gills or lungs. If these be not moistened with fresh or salt water then we are met with two alternatives —
either tho interchange of gases must take place through a membrane which after a time becomes dry, or the
membrane must be kept moist by an exudation from the animal. Certain observations on tlie genus Cocnohita
lead the writer to suppose that the latter is probably the case.
586 ON THE YOUNG OF THE ROBBER CRAB.
taken at night by torchlight running about the rocks at the brink of the sea, the
females having brown eggs attached to the abdominal limbs in large masses.
The copulation is at present unknown. There are no special organs to this end in
the male, which is withcjut abdominal limbs save on the sixth segment, and has no
such structure on the co.xopodite of the last leg as is found, for instance, in Coenobita
perlatus. This joint is, however, somewhat modified, being produced into a conical
process and connected with its fellow across the middle line. The process of coition
probably takes place inland, as females with eggs are fnund at long distances from
the coast (22). •
The eggs are borne in grape-like bunches attached to the long hairs on the great
biramous limbs of the second, third, and fourth abdominal segments. Each hair forms
the stalk of a bunch, and by displacing the eggs it can be seen that they are not
arranged evenly, but are in clumps of about half-a-dozen at intervals along the hair.
The mode of attachment is that commonly found among Decapods, namely by pro-
longation of the outer shell into a hollow stalk. The shape is ellipsoidal, and the
dimensions 8 mm. by •? mm. (in spirit). When the young begin to hatch the mother
washes them off into the water.
The embryonic skin must be very early lost, for, out of a large number of ne\vl\--
hatched young, every one had already thrown it off. Luckily the material on which
the present account is based, contained, among a number of unripe eggs, one which was
nearly read\- to hatch. By opening this egg a specimen still in the first skin was
gotten. In this it could be seen that the shape of the skin resembled on the whole
that shown for other pagurine zoaeas b\- G. 0. Sars (27). Further allusion will be made
to it in describing the telson of the free larva.
The first zoaea (Fig. 1), which is in no way remarkable in general appearance, has a
total length of rather over 3^ mm., more accurate measurement being impossible in the
Fig. 1.
Zoaea of the Robber Crab. The carapace has lifted somewhat from the body.
preserved specimens owing to the lifting of the carapace from the body. The carapace
is like that of other pagurine zoaeas in its principal features. There is a moderately
long rostrum, broad at the base but narrowing rapidly towards the free end. With this
exception the carapace is without spines. The hinder edge is hollowed and at each side
of the indentation is a rounded side-lobe. The length in the middle line, measured from
the tip of the rostrum to the hinder edge, varies slightly but is always rather under
li mm.
ox THE YOUNG OF THE ROBBER CRAB. 587
The abdomen consists of six segments, of which the last no doubt represents the
fused sixth segment and telson, as in Eupagurus, etc. (23, 27, 28). The segments are
Fig. 2.
Second antenna.
unarmed, save that the fifth bears a pair of stout teeth, one on each side, at the
hinder end. The telson (Fig. 8) has the usual fan shape with a shallow median notch as
in Eupagurus. On each side of the notch are five long bristles, equal in length and
Fig. 3.
Right mandible, seen under the microscope as a transparent object.
all moveably articulated. These bristles, like many of those on the limbs, bear a
thick fringe of very fine hairs. At the outer angle on each side is a stout immoveable
tooth. This arrangement, which closely resembles that figui-ed by MuUer (25) for an
unnamed species of Hermit Crab, differs from that in Eupagurus in having the bristles
of equal length and all moveably articulated, and from Spiropagurus in the median notch
Fig. 4.
First maxilla.
and the equality of the bristles. The embryonic skin fits the whole structure like a
glove, having processes corresponding to the teeth and bristles, but no supernumerary
w. V. 78
588
ON THE YOUXG OF THE ROBBEK CRAB.
ones. These processes are not fringed. It will be seen that Mayer's primitive form
of telson does not occur in Birgus (26).
On the other hand, the condition found in sundry other Macrura Anomala
(Eupagurm, Spiropagia-us, Galathea) in which the fourth bristle is longer than the rest,
fore-shadowing the arrangement in the crabs where this becomes the end of the
telson-fork, is also wanting in both the embryonic and larval telson of the present
genus.
Fig. .5.
Second maxilla.
The limbs are eight in number, rudimentary thii-d maxillipeds being present. The
first antennae are simple structui-es, unjoin ted and bearing hairs at the tip. The second
antennae (Fig. 2) consist of a basal joint prolonged without articulation into the endopo-
dite, and bearing as exopodite a broad scale resembling that of Spiropagurihs (27) with a
tooth at the outer ahgle and feathered bristles on the inner edge. The usual spine
at the base of the exopodite is also present on the basal joint of the limb.
Fig. C.
First maxiUiped.
The mandibles (Fig. 3) are somewhat highly developed, that on the right more so than
the left one. The former has a strong apical tooth, a cutting edge, and a complicated
Fig. 7.
Second maxilliped.
molar-like process. The latter two divisions are less strongly marked in the left
ON THE YOUNG OF THE ROBBER CRAB.
589
mandible. There is no palp. The first maxillae (Fig. 4) are much like those of Spiro-
pagurus. The second maxillae (fig. 5) bear six unusually distinct lobes. The three pairs
of maxillipeds show a strong resemblance to those of Eupagurus and Spii-opagui'us as
described by G. O. Sars (27).
The upper and lower lips are also much like those of the above two genera.
The jjaired eyes present that elongation of the crystalline cones of the hinder
half which is seen in other pagurines and in Thalassinidea (28). I was not able to
discover a " nauplius eye," but conclude that its absence is due to the state of
preservation of the specimens.
Fig. 8.
Telson.
If we now attempt to summarise in a few words the affinities of the zoaea of
Birgus, it may be said that the characteristics are those of the Anomalous Macrura
in general and of the Pagurinea in particular, that in most points it resembles Spiro-
pagurus rather than Eupagurus, but in the form of the telson inclines somewhat to
the latter genus, and lastly that its special peculiarity lies in an absence of spines and
teeth from parts of the body where they are found in allied forms and in a certain
simplicity of form throughout the whole organism.
The first larva of the Robber Crab thus proving to be a very ordinary zoaea, it
would seem probable that the further course of development presents no unusual feature
till the final stages are reached in which the peculiar adult habitus is assumed. This
is a point on which information is still desirable, as also on the details of the period of
transition from a sea life to one on land.
In ending, the writer must expre.ss his sense of the great obligation under which
Dr Willey has laid him by providing the material on which the above remarks are
based, consisting of a number of larvae and eggs and some valuable notes.
78—2
590 ON THE YOUNG OF THE ROBBER CRAB.
In view of the interesting nature of the habits of this crab and the amount of dis-
cussion to which they have given rise, a somewhat full list of works referring to the
natural history of the creature may be useful. Besides those mentioned below, several other
authors make passing reference to the habits of the robber crab.
I. THE ROBBER CRAB.
1. RuMPHius, "Amboinsche Rariteit-kamer," i., p. 7 (1705).
2. Linnaeus, "Systema Naturae," 12th ed., i., 2, p. 1049 (1767).
3. Herbst, " Xaturgeschichte der Krabben u. Krebse," ii., p. 34 (1791).
4. QuoY ET Gaimard, "Voyage de I'Uranie," Zool., p. 536 (1824).
5. Montgomery, "Voyages of Tyerman and Bennett," ii., p. 33 (1831).
6. Note in Proc. Zool. Soc, 1832, p. 17 (1832).
7. Dana, "U.S. Exploring Expedition," Crustacea, ii., p. 435 (1852).
8. Darwin, "Voyage of the Beagle," p. 462 (1870).
9. Von "Willemoes-Suhm, "Briefe von der Challenger-Expedition," Zeit. f. wiss. Zool., .xxvi.,
p. Lxxiii. (1875).
10. Bodd.\m-\Vhetham, "Pearls of the Pacific," p. 237 (1876).
11. Street.s, "Contributions to the Natural History of the Hawaiian and Fanning Islands,"
Bull. U.S. Mus., vii., p. 119 (1877).
12. Semper, " Ueber die Lunge von Birgus latro," Zeit. f. wiss. Zool., xxx., p. 282 (1878).
13. Semper, "Animal life," English Ed., pp. -5, 193, 449 (1881).
14. Guppv, "Note on the Coconut-eating habit of the Birgus in the Solomon Group," Proc.
Linn. Soc. N.S.W., vii., 4, p. 661 (1883).
15. Henderson, "Challenger" Report, Zoology, xxvii., Anomura, p. 50 (1888) [contains a
list of references, mostly systematic, up to 1888].
16. BouviER, "Le Crabe des Cocotiers ou Birgus larron," Naturaliste, xiii., 98, p. 81 (1891).
17. MosELEY, "Notes by a Naturalist on H.M.S. Challenger," p. 348 (1892).
18. Stebbing, "A History of Crustacea," p. 156 (1893).
19. Borrad.aile, "On some Crustaceans from the South Pacific, ii. Macrura anomala," Proc.
Zool. Soc, 1898, p. 457 (1898).
20. Borr-\daile, "A Note on the Hatching-stage of the Pagurine Laud-Crabs," Proc. Zool.
Soc, 1899, p. 937 (1900).
21. Ortmann, Bronn's " Thierreich," v., Crustacea, 2, p. 1235 (1900).
22. Andrews, "A Monograph of Christmas Island," p. 164 (1900).
II. THE LARVAE OF THE PAGURINEA.
23. R.\THKE, " Zur Eutwicklungsgeschichte der Decapoden," Wiegmann's Archiv. vi. (1840),
Transl. Mag. Nat. Hist, vi., p. 263 (1840).
24. Bate, " Carciuological Gleanings, No. iv.," Ann. Mag. Nat. Hist. (4) ii. (1868).
25. MiJLLER, "Facts and Arguments for Darwin," English Ed., p. 54 (1869).
26. Mayer, "Zur Eutwicklungsgeschichte der Decapoden," Zeit. f. Naturwiss. xi. (1877).
27. S.\RS, "Bidrag til kundskaben om Decapodernes Forvandlinger, ii.," Arch. Math, og
Naturvid. .xiii., p. 133 (1889).
28. Korschelt and Heider, "Te.xt-book of the Embryology of Invertebrates," English Ed.,
II., p. 286 (1899).
ANATOMY OF NEOHELIA PORCELLANA (MOSELEY).
By EDITH M. PRATT, M.Sc,
The Owens College, Manchester.
Plates LXII. and LXIII.
Neohelia is a rare geuus belonging to the imperforate Madrepores. Duncan in his
Revision of the Madreporaria places it with two other genera — Baryhelia and Dihlasus
in the alliance Baryhelioida in the family Oculinidae. It is interesting in that it is
the onlj' living representative of the " alliance," the other two genera having only been
found fossil — Baryhelia in the cretaceous deposits in England and Europe, Diblasus
in the upper cretaceous deposits in England. Baryhelia forms a massive colony, and
Diblasus is encrusting, the colony being irregular in shape.
The "alliance" that comes next to the Baryhelioida is the Lophohelioida, which
has at least two recent genera. It is clear from Fowler's description of Lophohelia
and Amphihelia that although there are certain general resemblances between the
two alliances as regards the anatomical characters of the soft parts, there is sufficient
difference to justify the retention of Duncan's classification.
The genus Neohelia contains the single species N. poixellana, which was first
dredged by the Challenger Expedition off Api Island, New Hebrides, at a depth of
63 fathoms. Three specimens each encrusting the stem of a Gorgonid were taken,
and the genus and species subsequently described by Moselej' in his Report on the
Challenger Corals, p. 176, PI. X. figs. 7, 7a, 18S1. His account however is restricted to
a description of skeletal structures.
In the collection of corals made by Dr Willey in the S. W. Pacific there
occurred a fragment of a colony of this species with the soft parts in a well preserved
condition.
A superficial examination revealed the following points of difference from the
publi-shed description of the species : —
1. All the Challenger specimens encrust Gorgonid stems. Our specimen is in
the form of an irregular hollow tube, and there is no evidence that this tube has
ever been occupied by a Gorgonid or other foreign support. 2. In his description,
592 ANATOIMY OF XEOHELIA PORCELLAXA (mOSELEy).
Moseley omits to mention certain apertures which are present in the peritheca'
(= coenenchyma) of a dried Challenger specimen now in the British Museum. These
apertures are larger and more numerous in our specimen which appears to be younger,
the peritheca is also less diffuse, and the costae more distinct, (compare Moseley,
Challenger, PI. X. figs. 7 and 7 a with figs. 1 and 2). 8. The hollow tube com-
prising our specimen is lined by a horny membrane which lies in contiguity with
the calcareous skeleton with which it conforms in outline and is interrupted in like
manner at the apertures (Fig. 1).
These were the chief points of interest together with an opportunity of ex-
amining the soft parts which led me to undertake the investigation of this species,
the results of which are now recorded.
Calcareous skeleton : —
In his description of the peritheca Moseley says, "The surface of the coenenchym
is marked all over by very slightly elevated rounded ridges which traverse it irregu-
larly, but with a general longitudinal direction, and are continuous at the margins
of the calicles with the costae." These ridges show more clearly in our specimen,
than in Moseley's, they are better seen after the soft parts have been removed by
immersion in a dilute solution of Caustic Potash followed by brushing gently. The
ridges are then seen to be studded with gi-anular protuberances. These however are
even better seen in a calcined specimen.
Septa. There are twenty Septa arranged in three cycles. While the number of
the Septa is constant in all the calicles which Moseley and I have examined, in some
cases, there is a tendency for them to lose their symmetry of arrangement (see
calicle S. Fig. 2). In Lophohelia prolifera (12), also a member of the Oculinidae,
there is a greater irregularity in the number and arrangement of the Septa. Fowler
remarks of this species that the total number of the Septa probably varies with
the age of the individual polyp.
Peritheca. The apertures in the peritheca are larger and more numerous in our
specimen than in Moseley's. In the latter they are so small, that their significance
was perhaps overlooked, but Moseley points out that as the colony covers the invested
^objects, the branches of the groiiS'ing coenenchym (peritheca) are soldered together.
The apertures referred to above and shown in Fig. 1, are gaps left between the
branches as they are soldered together, and probably diminish in size as the colony
grows older.
The Horny Membrane. In Moseley's. specimen which is dried, it is difficult
to ascertain if a horny membrane exists between the Gorgonia stem which the colony
encrusts and the calcareous skeleton, but as a horny substance of a similar nature
to that in our specimen forms a ring round one of tlie apertures, there is reason
to believe that the horny membrane is present generally in the genus.
1 Peritheca— teim originally used by Milue-Edwards and Haime, aud defined by Stanley Gardiner (24),
p. 361. 'The Peritheca is that part of the corallum of colonial Madicporaria, which is deposited outside
and subsequently to the theca.'
ANATOMY OF NEOHELIA PORCELLANA (mOSELEV). 593
Coenosarc. The coenosarc extends over the surface of the corallum from polyp
to polyp but has the same poral interruptions as the peritheca. The coenosarc is
traversed by numerous branching and anastomising Endodermal canals which are con-
tinuous with the coelentera of the polyps. One canal is given off from each inter-
mesenterial space in a manner similar to that of Stylophora described by G. von
Koch (19), by Stanlej' Gardiner for Coenopsanimia (24), by Fowler for Rhudopsuniniia (10).
Neohelia being an Im]ierforate madrepore, these canals are necessarily confined to the
surface of the colony.
There is nothing worthy of note about the general structure of the coenosarc, but
in the neighbourhood of the apertures in the wall of the colony, it is covered by a
.specialized epithelium which completely surrounds each aperture (Figs. .5 and 6, Ep. 1.).
This epithelial layer is seen to contain : —
1. ^suraerous large nematocysts with long si^irally-barbed threads similar to those
occurring on the acontia (Fig. 7).
2. Numerous gland cells, some of which, when stained with iron haematoxylin,
have a reticular appearance, while others contain granules (Fig. 6).
3. Interstitial cells of two kinds (i) of a long narrow columnar tj'pe, lying between
the nematocysts and the gland cells, (ii) more or less triangular cells which fill up
the spaces, below the nematocysts and gland cells and above the Mesogloea (Mg).
Below the epithelium and lying in the Mesogloea is an endodermal ring canal
which encircles the epithelium surrounding each aperture, this canal is lined by some-
what cubical cells with large nuclei, and sends off ramifications in the coenosarc which
comnuuiicate with the coelentera of the polyps (Fig. 5).
Neohelia is like Madracis in that it has a well-developed peritheca, it is also
like it in the fact that the peripheral lamellae of the mesenteries extend into the
coenosarc covering the peritheca. It is supposed by Fowler in Madracis that with the
growth of the peritheca resulting in the formation of echinulations, the coelenteron
between the lamellae becomes broken up into canals which form the only communication
between the polyp cavities. The canals in the coenosarc of Neohelia may be accounted
for in a similar way.
The important features noticed in the colony- are : —
1. A Horny-membrane lines the hollow tube which forms the colony.
2. There are certain apertures in the wall of the tube extending through coenosarc,
peritheca, and horny-membrane. As the colony grows older the apertures become smaller.
3. An " epithelial layer " and an endodermal ring canal encircles each aperture,
the latter communicating with the coelentera of the polyps. It is probable that the
" epithelial layer " encircling each aperture is of the same nature as the free growing
edge of the colony, (which unfortunately has been broken off), in having the power of
growth, the growing coenosarc secreting horny membi-ane and calcareous skeleton, and
thus filling up the apertures.
594 ANATOMY OF NEOHELIA POECELLAXA (mOSELEy).
There is really very little doubt that the horny substance is secreted by the
Neohelia itself, for it is in a thin continuous layer, and conforms in outline with the
calcareous skeleton, and, like it, is interrupted at the apertures in the wall of the
tube. If the horny membrane were secreted by another animal, we should not expect
to find these poral interruptions, and we should expect to find some trace of the
animal which formed it.
In the Alcyonaria and among the Zoantharia — the Actiniaria and the Antipa-
tharia — there are many instances of the secretion of a horny substance, therefore, it is
not improbable that such may occur among the Madreporaria. The layer of horn lies
in contiguit}" with the calcareous skeleton, and I have not observed any aggregation
of cells between the homy substance and the skeleton b}" which the former could be
secreted, but it is possible that it may be secreted by the epithelial layer, in a
manner similar to the secretion of the calcareous Epitheca by the Randplatte in a
form like Caryophyllia.
It is probable that the young coral polyp after settling down, has the power of
secreting, not onlj' a skeleton of lime but also a homy substance as in the Gorgonids.
As the 3"oung coral grows, it tends to encrust the object to which it has affixed
itself. This it would do in a more or less irregular manner, leaving gaps here and
there, — the apertures to which I have referred. As growth proceeds, the holes
become filled in until they are finally obliterated. (Compare Moseley's figures ^vith
Fig. 1.)
Among the Alcyonaria, there are two nearly allied genera, Suheria and Soleno-
caidon (2.5) belonging to the family Briareidae which show certain similarities as
regards their manner of growth with the genus Neohelia. Wright and Studer say of
Suheria gentJm, " The colony forms at first an encrusting surface over the long denuded
stems of a Gorc/onia. On reaching the termination of the foreign axis, and on several
occasions at intervals along its length, little upright stems arise, sometimes simple, but
sometimes branched."
In the case of Neohelia, I have seen two specimens, one (Moseley's) encrusting
a Gorgonid stem, the other erect and free but the base missing. It is probable that
the colony begins by encrusting a Gorgonid stem and afterwards sends off free branches.
Our specimen is probably one of the free branches.
The genus Solenocaulon (25 and 20) allied to Suberiu is characterised by the
tubular form of the axis which is perforated as in Neohelia, the tubes being formed
of a thin coriaceous substance. Gray says of it, " The coral cannot be considered as
a solid stem becoming hollow, as the last formed (younger) parts at the ends of the
branches are in the form of a foliaceous expansion, which gradually folds up together
on itself, coalesces and forms a tube nearly of the same diameter as the main stem.
The large apertures which occur in the stem and base of the branches, and communi-
cate with the central cavity are the parts of the expanded lamina which have not
been closed in when the other portions of the tube were formed." It is very probable
that the tube of Neohelia is formed in like manner, if so, then the apertures are
analogous, i.e. the holes in the wall of the tube of Neohelia are spaces which have
not been filled in by the peritheca.
ANATOMY OF NEOHELIA PORCELLANA (mOSELEY). 595
The species is probably monoecious, our specimen being a male colony.
Polyp. On measuring several polyps after the hard parts had been removed by
decalciticatiou, they were found to be 1*75 — IS mm. from top to bottom, and 1 — Vo mm.
in diameter. The polyps are closed at the base, and their coelentera are connected
by the system of canals which lie in the coeuosarc. There are no zooxanthellae
present in the tissues of this coral ; in this respect Neohelia resembles Coenop-
sammia (24).
Tentacles. The tentacles, twenty in number, are arranged in a single ring round the
periphery of the mouth disc, and correspond in position with the septa (Fig. 3). There
are five primary, five secondary and ten tertiary tentacles. They are all simple, short
and thick, many of them are bent over towards the mouth, while some of them are
partially invaginated, it is possible that they can be wholly invaginated. The nemato-
cysts, "02 mm. in length when unexploded, are of the common Actinian tj'pe, and are
more or less uniformly distributed over the surface of the tentacles in a manner
similai- to that described by Fowler in Turhinaria (12); they are perhaps more numerous
at the tip, but they are not arranged in knobs or batteries as in Lophohelia,
Amphihelia, Madracis, etc. Gland cells are numerous between the ncmatocysts and
interstitial cells which compose the free edge of a tentacle.
The Mouth Disc contains numerous large nucleated gland cells (Fig. 10). On
staining witii iron haematoxylin, some of the cells are seen to be filled with a deeply-
staining, hduiogeneous substance resembling mucous. In others the deeply-staining
substance forms a reticulum, while in a few cases the cells seem to be tilled with
a granular substance (Fig. 10 Gl). Nematocysts like those in the tentacles are numerous
on the mouth disc.
The Mouth is oval in cross section, with one large siphonoglyph and one small
one.
The walls of the Stomodaeum are nmch convoluted, probably due to contraction,
the Ectoderm, the histology of which could not be made out, is separated by a con-
siderable space from the Endoderm and Mesogloea ; this is probably due to the same
cause.
Mesenteries. There are twenty mesenteries arranged in three cycles, five
primary, tive secondary, and ten tertiary mesenteries. Only the primary and secondary
mesenteries support the stomodaeum, the tertiary mesenteries extend only for a very
short distance into the body cavity. There are two pairs of directive mesenteries. The
ten principal mesenteries bear mesenterial filaments and acontia. Any of these mesen-
teries may bear generative organs, in one case they are borne on the directives. In
most of the polyps I have examined only one or two of the mesenteries bear gene-
rative organs, but in a few cases they occur in five or six of the principal mesenteries
in a polyp. The tertiary mesenteries have no mesenterial filaments or acontia, and
they do not bear generative organs.
Upon examination with iiigh powers of .sections staimcl with burax carniiin' I have
seen at th(! insertion of the mesenteries certain structures similar to those described
w. V. 79
596 ANATOMY OF NEOHEI.IA PORCELLAXA (mOSELEY).
by Bourne (4) in Mussa and called by him " desmocytes." Owing to their minute
size in this foi-m I have not been able to confirm Bourne's results as regards other
details of these interesting structures.
Mesenterial filaments as stated above occur only on the primary and secondary
mesenteries. In transverse section they are seen to be three lobed. Gland cells are
numerous, but no nematocysts have been seen on them. In this respect Neohelia
agrees with Flahellum patagonichum, of which Fowler (10, p. 16) says:
" Nematocysts do not occur apparently in- the true mesenterial filament, but only
on that portion of it which is continued on to the contorted lamellae, which I regard,
in part at least, as equivalent to the acontia of Actiniae."
Acontia. The lower portion of the coelenteron is occupied by numerous groups
of structures coiled together. In some cases they are protruded through the anterior
body wall outside the crown of tentacles. In section they are then seen to be more
or less rounded structures occurring in groups of four or five arising from a common
stem which is given off from the free edge of a mesenter}^ (Figs. 8 and 3). These
acontia, for such I believe them to be, are armed with numerous large nematocysts
(Fig. 7) which, when exploded, are seen to be provided with a formidable spirally-
barbed thread of great length. Similar nematocysts in an uuexploded condition were
observed on the epithelium bordering the apertures in the tubular wall of the colony.
Length of unesploded nematocyst "038 mm. Gland cells are numerous round the
periphery of the acontia, the internal portion of an acontium is made up of spindle-
shaped interstitial cells, developing gland cells and nematocysts. The stem supporting
the acontium is composed of an axis of mesogloea which extends into and branches
in the acontium in a manner similar to that described by Bourne in acontium of
Fungia (3). The mesogloeal axis is covered by a single layer of eudodermal cells.
Of Flahellum patagonichum (Moseley) Fowler says: "What Moseley has termed 'the
contorted mesenterial filaments,' a mass of coils lying on the side of the mesenteries,
appear to me after careful investigation to be, in part at least, organs corresponding
to the acontia of Actineae, namely, long lamellar offshoots of the fi-ee edge of the
mesentery with one edge thickened to correspond to the mesenterial filament, and
charged with large nematocysts. They protrude in some instances through definite
openings in the mouth disc. Their exact origin fi-om, and relation to the mesenteries
I have not been able to detect owing to the brittle condition of the specimens."
There seems to me to be little doubu that these structures described by Fowler
in Flahellum are very similar to the acontia described by Bourne in Fungia, and to
those occurring in Neohelia. In the latter, however, I have observed no instance in
which the acontia are thrust through apertures in the mouth disc, in every case,
which I have examined, they are everted through the anterior body-wall outside the
crown of tentacles.
ANATOMY OF NEOHELIA PORCELLANA (mOSELEY).
597
Rklations ok Neoiiema.
Our knowledge of the anatomy "I' the corals allied to Neohelia is still very
incoini)lete, hut the foilowiiic; tables may be some guide to us in the discussion of its
artinities.
A comparison of the hard parts of Neohelia with the allied fossil genera
Buri/helia (20) and Diblusus (17) is given in the table subjoined.
Unfortunately the description of Barijhelia from which this comparison is drawii
up by Milne-Edwards and Haime is meagre and unaccompanied by figures.
Neohelia
Barybelia
Diblasus
Colony
Partly encrusting
„ erect
Mas.sive
Encrusting and
irregular in shape
Peritheca
Abundant, diffuse
costiilate iind gi-anulated
Moderately developed
smooth or finely gniuulatod
Moderately developed
co.stulatc
Calices
Projecting, small
diameter 1 5 ni.m.
height 2 m.m.
Slightly projecting
or not, .Small
Projecting, irregular in
size. Average diameter
5 m.m.
Septa
•20 in 3 cycles
unequal, "ranular, often
fiused together
Few, entire
thick, short
3 cycles, unequal, dentate,
crowded ami granular.
Larger liecome fused
Columella
Alwent
Absent
Rudimentary, formed by
fusion of liirger septa
Gemination
Irregularly
dicliotomoiLs
Marginal and
intercalicinal
Time
Recent
Foeail
Cretaceous
Fossil
UpiJer cretaceous
Fmm this comparison which, it is needless to state, is very incomplete, it would
appear that Diblasus is more nearly related to Neohelin than is liari/fielid. When
further investigation of the latter is made a nearer relationship between the two genera
may be found to exist.
Neohelia differs chiefly from DiblasKs,
1 In lh:it it may form a liuljuw tul)e.
•J. In till' presence of gaps in the peritheca.
70—2
598
ANATOMY OF NEOHELIA PORCELLANA (mOSELEy).
3. In the presence of a horny membrane, but as this is fairly delicate ib
could scarcely be expected to be present in the fossil form.
4. In the absence of a columella which is rudimentary in Diblasus.
A comparison of Neohelia with other members of the Oenlinidae.
Tentacles
Mesenteries
Neohelia
20 in number
(5 primary, 5 secondary, 10 tertiary)
Simple, short and thick with nemato-
cysts uniformly distributed over the
surface, not in knobs or batteries.
Correspond in pcsition with the .septa
20 in number
(5 primary, 5 secondary, 10 tertiary)
Primaries and secondaries bear me-
senterial filaments and generative
organs and acoutia.
2 pairs of directives.
Generative organs Actinian
Lophohelia
Number (?)i
Knobbed; each knob is a battery of
nematocysts. Con-espond in position
with .septa
Vary in number in different polj-ps.
No directives.
Amphihelia
Number (?)
Covered with batteries of nemato-
cysts. Correspond in position with
septa
12 — 14 pairs normal.
2 pairs directives.
Generative organs Actinian
Madracis
Number (?)
Simple, each tipped with a single bat-
tery of nematocysts. Apparently
correspond in position with septa
8 pairs.
No differentiation of particular me-
senteries recognisable. All extend to
about the same dejjth in polyp cavity.
2 pairs directives
Neohelia does not appear to be nearly related to any member of the Oculinidae
with which I have compared it, so it is necessary that, at present, it should be
placed as Duncan puts it in a separate alliance.
The Research in connection with this paper has been done iu the Zoological
Laboratories of the Owens College, Manchester, under the supervision of Professor
Hickson, to whom I am greatly indebted for much assistance and advice.
1 Fowler (12), in his description of Lophohelia remarks that the tentacles are arranged one over every
septum. Septa vary in number in different Individuals, so that one would suppose that the number of
tentacles also varies.
ANATOMY OF NEOHELIA PORCELLANA (mOSELEY). 599
1. AsuwoRTH, J. H. Stomodaeum, mesenterial filaments and Endodemi of Xenia. Proc.
Zool. Soc. Lond., lxiii., p. 443.
2. AsHWORTH, J. H. Xeniidae. Willey's Zool. Results. Pt iv., 1900, p. 509, Pis. LII., LIII.
3. Bourne, G. C. Anatomy Madrepor. Coral Fungia. Quart. Journ. Mic. Sc. Vol. xxvii.,
pt 3, N.S.
4. Bourne, G. C. Calcareous skeleton of the Anthozoa. Quart. Journ. Mic. Sc. Vol. xli.,
N.S., Jan. 1899.
5. Dana. Zoophytes.
6. Duncan. Revision of the Madreporaria.
7. Duncan. Deep Sea Corals. Nature, Vol. 30, p. 464.
8. Duncan. Diblasus. British Fossil Corals. Supp. to Pal. Soc, Pt ii.. No. 1, 1869,
p. 14, PI. II., figs. 1—11.
9. Duerdin. Relations of certain Stichodactylinae to the Madreporaria. Journ. Linn. Soc.
Zool. XXVI.
10. Fowler, G. H. Anatomy of the Madreporaria.
I. Flabelluni, Rhodopsammia. Quart. Journ. Mic. Sc, Vol. xxv.
11. Fowler, G. H. II. Madrepora. Quart. Journ. Mic Sc, xxvii.
12. Fowler, G. H. III. Turbinaria, Lophohelia, Seriatopora, Pocillopora. Quart. Journ. Mic.
Sc, Vol. xxviii.
13. Fowler, G. H. IV. Madracis, Amphihelia, Stephanophyllia, Sphenotrochus, Stephanaria,
Pocillopora, Seriatopora. Quart. Journ. Mic. Sc, Vol. xxviii.
14. Fowler, G. H. V. Duncania, Madrepora, Galaxea, Heteropsammia, Bathyactis. Quart.
Journ. Mic. Sc, xxx.
15. Gray, J. E. Solenocaulon. Zool. Soc, Feb. 1862. Abst. in Annals and Mag. Nat.
Hi,st., 1862.
16. Hertwig, O. & R. Die Actinien. Jena, 1879.
17. Hickson, S. J. Anatomy Alcyonium digitatum. Quart. Journ. Mic. Sc, Vol. xxxvii.,
p. 343, 189.5.
18. G. von Koch. Mitth. ii Colenteraten. Jena Zeitsch. Bd. xi. (10) Abst. in Fowler.
Anat. Madreporaria i.
19. G. von Koch. Mittheilungen iiber das Kalkskelet der Madreporarien. Morph. Jahrb.
VIII., p. 85. Abst. in Journ. Mic. Sc, N.S., ii., p. 795.
20. KuKENTHAL. Solenocaulon Ergebnisse. Bd. xxii., pp. 149 — 161. Tafs. ix. Figs. 3, 4.
21. Milne-Edwards and Haime. Hist. Nat. des Coralliaires. Paris, 1860.
22. MosELEY. Report on the Challenger Corals. 1881. Challenger Reports, Vol. ii.
23. Ogilvie (Miss), M. M. Microscopic and Systematic Study of the Madreporarian types of
corals. Phil. Trans. CLXXXVii., 1896.
24. Stanley Gardiner, J. Anatomy of a supposed new species of Coenopsamniia from I^ifu.
Willey's Zool. Results. Pt iv.. May 1900. PI. xxxiv., Figs. 12 and 13.
25. Wright and Studer. Suberia and Solenocaulon. Report on Alcyonaria. Challenger,
Pt Lxiv. p. 163.
600 ANATOilY OF NEOHELIA PORCELLANA (mOSELEY).
DESCRIPTION OF PLATES LXII. AND LXIII.
Fig. 1. View of a portion of the colony magnified about four and a half diameters.
The colony forms an irregular hollow tube, the horny membrane lining the tube is sho^vTi
at the broken ends. The branches on the right side show a tendency to become fused
(this tendency is more clearly marked in Moseley's specimen). The granular ridges more or
less continuous with the costae are shown as dotted lines. (Del. E. R. Dust.)
G.R. granular ridges.
H.M. horny membrane.
P. apertures in tube.
Fig. 2. View of a portion of the skeleton after the soft parts have been removed —
by immersion in a solution of caustic potash followed by careful brushing.
The 20 septa are arranged in three cycles the primary and secondary of 5 each, and
the tertiary in a cycle of 10.
The costae are more ur less continuous with the septa and bear granular echinulations.
There is no columella, but the septa become fused at the bottom of the calicle forming a
wall which shuts ofi" the polyp from the remaining portion of the colony.
1. Primary septa.
2. Secondary septa.
3. Tertiary septa.
C. Costae.
G.E. Granular echinulations.
.S'. Calicle showing irregularity in arrangement of septa.
Del. E. R. Dust.
Fig. 3. Longitudinal section through a polyp. About 70 diameters. The section passes
through a tentacle on the left side and between two tentacles, i.e. through a mesentery on
the right side. Tiie dotted shaded portion represents the calcareous skeleton, and the
ectodermal portion of the stomodaeum is indicated by dark shading.
Ac. Single acontium with stem much contracted.
Can. Canal in coenosarc, its communication with the coelenteron of pol}^ is
not shown.
Ca. a. Calcareous skeleton.
Cy. Calycoblasts.
Uc. Ectoderm.
£n. Endoderm.
Mff. Mesogloea.
31. Mesentery.
WiLbEY. Zoological Resi;;
HM %Vj.
Can
CaS
■£n
ANATOMY OF NEOHELIA PORCELLANA (mOSELEY). 601
M.f. Mesenterial filament with gland cells {yl. c).
N. Nematocysts on acontiuin.
St. Stomodaeum.
T. Tentacle.
Tes. Testis.
Fig. 4. T.S. through a single calicle showing the distribution of hard and soft parts.
About 60 diameters. The dotted portion represents the calcareous skeleton.
The primary tentacles correspond in position with the primary septa, the secondary,
with the secondary septa, and so on.
In the coenosarc outside the polyp peripheral lamella enclosing portions of the coelenteron
and forming a portion of the Endodermal canal system, are seen lying in the mesogloea.
(Caw.)
Ca. Calcareous skeleton.
Cy. Position of calycoblasts.
Ec. Ectoderm.
En. Endoderm.
Mg. Mesogloea.
Mes. Mesentery.
P.S. Primary Septum.
S.S. Secondary Septum.
T.S. Tertiary Septum.
P.T. Primary Tentacle.
S.T. Secondary Tentacle.
T.T. Tertiaiy.
Fig. 5. Surface view of a portion of coenosarc in the neighbourhood of an aperture
in the wall of colony. The aperture is surrounded by a specialized epithelial layer, .£;;. X.,
which is encircled by an Endodermal Ring Canal, R. C. ; this gives off numerous branches
which ramify and anastomose in the coenosarc and connect the coelentera of the polj'ps.
(Cam. Luc. x 43).
A. Aperture.
Can. Canals.
Coen. Coenosarc.
Fig. 6. Section through Specialized Epithelium and Endodennal ring canal surrounding
hole in wall of colony (x 660. Iron Haematoxylin).
The Epithelium contains : 1. Nematocysts similar to those on acontia. 2. Gland cells.
3. Interstitial cells.
En. Endoderm.
GL Gland Cells.
In.C. Interstitial cells.
Mg. Mesogloea.
#. Unexploded Nematocysts.
R.C. Ring Canal.
Fig. 7. A single exploded Nematocyst of Acontium showing spirally barbed thread.
x870.
602 ANATOMY OF NEOHELIA PORCELLANA (mOSELEY).
Fig. 8. Longitudinal Section through a group of Acontia which are thrust outside the
body wall of pol}'p, outside the radius of the tentacles {Cam. luc. x 408), Semi-diagram.
Only one Acontium is drawn in detail. Round the periphery are seen the exploded
nematocysts and gland cells, the interior is filled with interstitial cells developing nemato-
cysts and gland cells and ramifications of the mesogloeal core of acontial stem.
Ac. Acontia.
£n. Endoderm.
Gl. Gland cells.
I.e. Interstitial cells.
Mg. Mesogloea.
If. Nematocysts.
Fig. 9. T. S. through a mesentery bearing Testis, x 300.
£n. Endoderm.
3Ig. Mesogloea.
M./. Mesenterial filament.
Te. Testis.
Fig. 10. Z. S. through a group of Gland Cells on Mouth-disc, x 700.
Un. Endoderm.
Gl. Gland cells.
In.C. Interstitial cells.
Mg. Mesogloea.
WiLLEY. Zoological Results
Plate LXI'
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C. W/'/son, Camirtctya,
(.-- KFOTTF]'/"-
ON A NEW BLIND SNAKE FROM LIFU, LOYALTY ISLANDS.
By G. a. BOULENGER, F.R.S.
With figures in the text.
In his excellent " Catalogue des Reptiles de la Nouvelle Cal^donie," published in
the Memoirs of the Linnean Society of Normandy, vol. xv. 1869, A. Bavay mentions
the common occurrence of a small Boid Snake, Enygrus Bibronii (now identified as
E. australis, Montrouzier) on Lifu at the point called Gatcha; and he observes that
it is the only terrestrial Snake found on that island, whilst none but marine forms
(HydropJiiuue) are known to occur in New Caledonia. Although, through one of those
absurd blunders which so frequently spoil the work of compilers, a poisonous land
Snake has appeared on lists of New Caledonian Reptiles', only Hydrophiines have to
the present day been seen in that archipelago. It is therefore very interesting now
to find the fauna of Lifu enriched by the discovery of a second land Snake. This
discovery we owe to Dr Willey, who brought home from that island a single specimen
of a Typhlops belonging to an undescribed species which it gives me great pleasure
to name
Typhlops Willeyi.
Diagnosis: Snout depressed, rounded, subacuminate, .strongly projecting; nostril
lateral, between two nasals, the anterior of which is in contact with the first and
second labials; nostril one-third the width of the head, not extending to the level of
the eyes ; praeocular present, much broader than the ocular, in contact with the second
and third labials; eyes distinct; prsefrontal, frontal, and interparietal not larger than
the scales on the body, supraocular a little larger, parietal twice as large; four upper
labials, third and fourth nearly equal and in contact with the ocular. Diameter of
body 32 times in the total length ; tail as long as broad, ending in a small spine.
22 scales round the body. Olive-brown above, yellowish beneath.
Total length I'J.J millimetres.
' See Trouessart, Hull. Sue. Zool. Francv, xxiii. 1898, p. 186.
w. V. 80
604
ON A NEW BLIND SNAKE FROM LIFU, LOYALTY ISLANDS.
The single specimen has been presented to the British Museum by Dr Willey.
By its technical characters, this species approaches nearest Typhlops acuticauda,
Peters, from the Pelew Islands, and T. aluensis, Blgr., from the Solomon Islands, both
of which differ, among other points, in the narrower praeocular shield and the much
more slender body.
Considering the general distribution of the Typhlops, the fact of a species
inhabiting the Loyalty Islands is far less remarkable than the total absence of repre-
sentatives in New Caledonia. In reality the herpetological fauna of the latter islands.
Profile, upper and lower views of head, and lower view of tail of Typhlops Willeyi.
by no means a poor one so far as Lizards are concerned, bears no kind of affinity
to that of the Loyalties. The high degree of specialization attained by some Geckos
of New Caledonia, viz. the curious species of the endemic genus Rhacodactylus, is even
borne out by the osteological structure, as I have recently ascertained the parietals to
be fused to a single bone in R. trachyrhynchus, thus presenting a remarkable
exception to a character which has been used to define families, or even " suborders "
among the Lacertilia. Another instance of the overrated importance attached to the
fusion of paired bones is to be found in the genus Varanus, where I have found the
nasal bones to be distinct in two adult specimens of V. salvator, one in the Museum
of the Royal College of Surgeons (as noted by me in 1891), the other in the
Collection of University College (as pointed out to me by Prof. Minchin), as well as
in the skeletons I have examined of V. prasinus and V. timorensis.
ON CRUSTACEA BROUGHT BY DR WILLEY FROM THE
SOUTH SEAS.
By the Rev. T. R. R. STEBBIXG, F.R.S.
With Plates LXIV.— LXXIV.
Already two reports have appeared on Dr Willey's extensive collection of Crustacea
from tropical waters and islands of the south-west Pacific. In 1898 a new species of
Caprellidae from Lifu was described by Dr Paul Mayer, and in 1899 Mr L. A. Borradaile,
after examining eighty-two species of Stomatopoda and Macrura, determined no less than
twenty of them to be forms new to science. In the lower groups the proportion of
new forms has proved to be even more considerable, so far at least as concerns the
specimens actually investigated. There is still a residuum of small creatures, of which
many but more probably few may prove to have been hitherto undescribed. For various
reasons these are omitted from the present report, the leading motives for this neglect
being that the report itself should not be indefinitely expanded or indefinitely delayed.
The species now recorded are forty-six in number, distributed over thirty-four genera,
of the Malacostraca, Entomostraca, and Thyrostraca. Of the genera eight are here for
the first time established, and of the species twenty-three are registered as new. Ex-
ceptional interest will be recognized as attaching to the Th\Tostracan genus which I have
named Koleolepas. Upon this I venture to quote from a private letter in which Dr Willey
modestly says, ' I have a foolish tendency to feel a trifle elated about Koleolepas n. g.
I imagined that it was not a very frequent occurrence for a new genus of Cirripede to
turn up, but this one struck me as being quite remarkable with its disc of attachment
and contractile cylindrical body as well as its peculiar paguroid habitat, although that is
very likely not constant.' Anchicaligus nautili (Willey) and Panaietis incamerata, both
from the pallial chambers of mollusca, are not undeserving of notice. From P. J. van
Beneden long ago to Mr Thomas Scott the other day, authors have called attention to
the large opportunity for finding crustacean parasites which almost the whole range of
the aquatic fauna provides. Of this it will be seen that Dr Willey has successfully
availed himself Neither in this nor in other respects has he been deterred by the
exigencies of his own special research from advancing collateral branches of knowledge.
Rather, he has utilized those exigencies for that very purpose, so that, at least in regard
to ' natural history.' he has earned a right to say, Scientiae nihil a me alienum puto.
80—2
606 ON CRUSTACEA BROUGHT BY DR WILLEY FROM THE SOUTH SEAS.
In regard to the nomenclature adopted in this report, I am bound to direct the
attention of critics and censors to an innovation, by which the term Cumacea, just as it
was at length becoming familiar, is displaced in favour of a new title. Of minor import-
ance is the defence here put forward for the use of the generic name Cubaris in prefer-
ence to that of Armadillo among the terrestrial isopoda.
The following list shows the species dealt with in the present report, with the
habitat from which each was obtained.
MALACOSTRACA.
Decapoda Macruea. Fam. Scyllaeidae.
Phyllosoma duperreyi Guerin. Milne Bay, New Guinea.
Sympoda. Fam. Nannastacidae.
Nannastacus ossiani n. sp. Barawon, Blanche Bay, New Britain.
Nannastacus (jeorgi n. sp. Barawon, Blanche Bay, New Britain.
Isopoda.
Fam. Tanaidae.
Leptochelia minuta Dana. Sandal Bay, Lifu, Loyalty Islands.
Leptochelia lifuensis n. sp. Lifu, Loyalty Islands ; and from sponge, Isle of Pines.
Fam. Anthueidae.
Apanthura sandalensis n. g. et sp. Sandal Bay, Lifu, Loyalty Islands.
Paranthura lifuensis n. sp. Sandal Bay, Lifu, Loyalty Islands.
Fam. Gnathiidae.
Onaihia aureola n. sp. Sandal Bay, Lifu, on sting-raJ^
Fam. Cirolanidae.
Cirulana pleonastica n. sp. Blanche Bay, New Britain.
Cirolana albicaudata n. sp. Barawon, Blanche Bay, New Britain.
Cirolana orientalis Dana. Conflict Islands, New Guinea.
Cirolana minuta Hansen. Lifu, Loyalty Islands.
Hansenolana anisopous n. g. et sp. Isle of Pines.
Fam. Alcieonidae.
Alciroiia imularis Hansen. Blanche Bay, New Britain.
Fam. Cymothoidae.
Anilocra dimidiata Bleeker. British New Guinea.
Renocila periophthalmi n. sp. Lifu, Loyalty Islands, on Periophthulmus.
Meinertia (jaudichaadii (Milne-Edwards). Panaieti, Louisiade Archipelago, New
Guinea.
Fam. Sphaeromidae.
Cilicaea tenuicaudata Haswell. Blanche Bav, New Britain.
on crustacea brought by dr willey from the south seas. 607
Fam. Ligiidae.
Ligia vitiensis Dana. Matadona, China Straits, British New Guinea.
Fam. Oxiscidae.
Philoscia gracilis Budde-Lund. Lifu, Loyalty Islands.
Philoscia truncata Dollfus. New Britain.
Philoscia lifuensis n. sp. Lifu, Loyalty Islands.
Paraphiloscia stenosoma n. g. et sp. New Britain.
Fam. Armadillidiidae.
Cubans transliccidus (Budde-Lund). Lifu, Loyalty Islands.
Citbai-is lifuensis n. sp. Lifu, Loyalty Islands.
Cubans doll/usi n. sp. Lifu, Loyalty Islands.
Cubans officinalis (Desmarest). Isle of Pines, S. of New Caledonia.
Cuharis lundi n. sp. New Britain.
Cubaris zebricolor n. sp. Lifu, Loyalty Islands.
Amphipoda.
Fam. Talitridae.
Parorchestia hawaiensis (Dana). Lifu, Loyalty Islands.
Fam. Rhabdosomidae.
Rhabdosoma whitei Bate. Blanche Bay, New Britain.
ENTOMOSTRACA.
Braxchiopoda.
Phyllocarida. Fam. Nebaliidae.
Nebalia bipes (0. Fabricius). Sandal Bay, Lifu, Loyalty Islands. Blanche Bay,
New Britain.
Ostracoda.
Myodocopa.
Fam. Asteropidae.
Asterope arthur-i n. sp. Blanche Bay, New Britain.
Fam. Cypridinidae.
Cypridina baravoni n. sp. Barawon, Blanche Bay, New Britain.
Copepoda semiparasitica.
Fam. Lichomolgidae.
Linchiomolgus caendeus n. g. et sp. China Straits, New Guinea.
608 ox crustacea brought by de ^villey from the south seas.
copepoda parasitica.
Fam. Caligidae.
Anchicaligus nautili (Willey). Lifu, Loyalt}- Islands, New Britain.
Gloiopotes hygomianus (Steenstrup and Llitken). Rubiana, New Georgia.
Fa5i. Dichelestiidae.
Bassettia congri n. g. et sp. Blanche Baj-, New Britain.
Pneudocycnus appendicidatus Heller. Uvea.
Panaietis incamerata n. g. et sp. Panaieti, Louisiade Archipelago, New Guinea.
THYROSTRACA.
(Gtj'Hpedia.)
Fam. Lepadidae.
Poecilasma vagans Aurivillius. Sandal Bay, Lifii, Loyalt}^ Islands, and New Britain.
Megcdasma striatum Hoek. Blanche Bay, New Britain.
O.vynaspis aurivillii n. sp. New Britain.
Concltoderma hunteri Darwin. Blanche Bay, New Britain.
Scalpellum sp. Blanche Bay, New Britain.
Koleolepas ivilleyi n. sp. Sandal Bay, Lifu, Loyalty Islands.
MALACOSTRACA.
Decapoda Macrura.
Fam. Sctllaridae.
Phyllosoma, Leach.
1818. Phyllosoma, Leach, Nat. Hist. App. to Tuckey's Narrative of an Expedition
to exjjlore the river Zaire, usually called the Congo, App. lY. p. 416.
182.5. Phyllosoma, Desmarest, Consid. ge'n. Crust., p. 2.53.
1833. Phyllosoma, Guerin, Magasin de Zool., el. 7 (unpaged).
1837. Phyllosoma, Milne-Edwards, Hist. Nat. Crust., vol. 2, p. 472.
1838. Phyllosoma, Gu^rin-Meneville, Voy. de la Coquille, Crustaces, p. 46.
1863. Phyllosoma, Claus, Zeitschr. wi'ss. Zool, vol. 13, pt 3, p. 422.
1873. Phyllosoma, Richters, Die Phyllosonien, Inaugui-al-Dissertation.
18S0. Phyllosoma, Boas, Vid. Selsk.'Skr., ser. 6, Nat. Afd. 1, p. 83 (61).
188S. Phyllosoma, Bate, Challenger Reports, vol. 24, Macrura, pp. 56, 89.
In the authorities above cited I think that almost everything will be found which
is at present known or surmised in regard to the larval forms composing this genus.
A single specimen in Dr Willey's collection must be identified with the species named
ON CRUSTACEA BROUGHT BY DR WILLEY FROM THE SOUTH SEAS. G09
below, which belongs to the Scyllarid group, with the second antennae broad in adult
and young. Mr Borradaile's report mentions Scyllarus sieboldi, de Haan, and Parihacus
antarcticus (Rumph), as obtained by Dr Willey, but these were from Lifu, Loyalty
Islands. Leach, who is known to be the author of appendix iv. to Tuckey's Narrative,
though his name is not expressly appended to it, introduces four species of Phyllosoma,
brevicoriie, laticorne, commune, and clavicorne, with distinguishing figures on plate 12.
He begins his account with the words, ' Phyllosoma, the most curious genus of Crustacea
that has yet been discovered.' Familiarity may lessen the curiosity of it, and the
o'eneric name is likely to disappear, when all the problems of affinity between these
larval forms and their parents have been solved. But the beauty of structure remains
exceptional, even now that many parallels are known to the otherwise amazing contrast
between the young and adult stages of an individual life.
Phyllosoma duperreyi, Gu^rin.
1833. Phyllosoma Duperreyi, Guerin, Magasin de zool., cl. 7 (unpaged), pi. 12.
1837. PJiyllosonia Duperreyi, Milne-Edwards, Hist. Nat. Crust., vol. 2, p. 485.
1S3S. Phyllosoma Duperreyi, Guerin-Meneville, Voy. de la Coquille, p. 40, pi. .5,
fig- 2.
1873. Pityllosoma Duperreyi, Richters, Die Phyllosomen, p. 17, in Zeitschr. wiss.
Zool, vol. 23, pi. 33, fig. 3.
It should be noticed that both Guerin (later Guerin-Meneville) in 1833 and Milne-
Edwards in 1837 give references to a work which itself contains a preface dated 1838,
Guerin citing " Voyage de Duperrey, Zool. t. II, p. 2, p. 46 ; pi. 5, fig. 2," and Milne-
Edwards citing " Voyage de la Coquille, Crust. PI. b, fig. 2." The explanation may
be that the plates were published before the text of Duperrey's voyage, and that
Guerin had in his hands the printed text of his report long before it was published.
The specimen he described and figured was taken at Port Jackson, and measured more
than 40 mm. in length by 30 mm. in breadth. Dr Willey 's specimen, from Milne Bay,
New Guinea, is 22 mm. long by 13 mm. wide. The specimen of another species, which
Guerin figures as Phylloso7na laticorne, Leach, is represented as three inches long by
nearly two inches wide, with a span between the extremities of its slender legs, when
the longest are fully extended, of twelve or thirteen inches.
SYMPODA.
1846. Cumacea, Kroyer, Naturhistorisk Tidsskrift, Ser. 2, vol. 2, ]). 203.
186."). Cumacea, G. 0. Sars, Vid.-Selsk. Forh. for 1864, Extract, pp. 1—83.
1893. Cumacea, Stebbing, History of Crustacea, Internat. Sci. Ser., vol. 74, pp. 8, 291.
1899. Cumacea, Sars, Crustacea of Norway, vol. 3, p. 1.
Since the important essay by Sars in 180-5 Kroyer's name for this order has been
accepted without denuir in a very large number of writings on the subject down to
the present date. Of all these works the most imjiortant is the third volume of the
610 ox CRUSTACEA BROUGHT BY DR WILLEY FROM THE SOUTH SEAS.
Crustacea of Norway, now on the eve of completion and entirely devoted to this group
under its long-received designation. It must be admitted that a change in the title
has become very inconvenient and is likely to be very unwelcome. Nevertheless, the
reasons for making it may be allowed to outweigh such objections.
When the principal divisions of the Crustacea are considered, whether they are
regarded as sub-classes or orders or sub-orders, it appears that a name founded upon
a particular genus is limited to the Cumacea, except in the case of the recently
separated and not universally accepted order or sub-order of the Tanaidacea. But the
Cumacea, if allowed to derive their ordinal name from a genus, should at least derive it
fi-om the earliest generic name within the group, which is Diastylis, established by Say
in 1818, not Cuina, due to Milne-Edwards in 1828. Illustrious as Hemi Milne-Edwards
personally was, there is no great reason for honouring him through the name of this
genus, which he persisted in regarding as of larval character. Moreover, the word
Guma itself is under a cloud. Scudder's Nomenclator Zoologicus, an easily accessible
work, gives from Agassiz, " Cuma Humph. Moll. 179-5. A." Humphrey's work is rare,
but Chenu, Manuel de Conchyliologie, vol. 1, p. 171, 1859, is still using his genus
for two species of Gasteropods, C. angidifera and C. kiosqmformis, of Duclos, heading
the description with the words, " SI" Genre. Cuma. Humphrey, 1797. Etabli aux depens
des pourpres." Lately with the friendlv assistance of Mr B. B. Woodward at the
British (Natural History) Museum I have been able to compare Scudder and Chenu
with the original authority for the name in question. The book in which it occurs,
bearing the title 'Museum Calonnianum,' was published anonymously in 1797, but is
known independent!}' of its title-page to have been by G. Humphrey. It contains at
p. 35, between the genera Fums and Mitra, 'Genus LX. Cuma — L'Onde — Wave,'
followed by 13 species, numbered from 645 to 657. The first of these is named
' Icterica,' but this being, like most of the others, unaccompanied by any description,
figure, or reference, must be regarded as a nomen nudum. On the other hand
' 646. Aulica ' is referred to ' Buccinum Aulicum Soland.,' and ' 647. Morio ' is referred
to 'Voluta Morio Linn. Buccinum Morio Soland.,' while to '650. Prismatica' there is
appended a short description by the author himself There is no angidifera or
kiosquiformis among the original species, so that Chenu's use of the genus for none
but those two is illegitimate. None the less it is evident that a molluscan genus
Cuma was established, in however poverty-stricken a manner, in 1797, and that the
crustacean genus which received this preoccupied name in 182s must fall back upon
some other designation. The result is that the displacement of Cuma, Milne-Edwards,
will lead to the reinstatement of Bodotria, Goodsir, and make the name Cumacea wholly
inappropriate and unmeaning.
The new title here offered is derived from the Greek a-v/j.Trov<;, av/j,'7roBo<;, meaning
' with the feet closed together.' This is sufficiently characteristic of the general appear-
ance, and the name has the advantage of agreeing in termination with the names of
the neighbouring groups, the Amphipoda and the Isopoda, which owe their titles m
like manner to what may be called an impressionist estimate of the limbs.
on crustacea brought uv uli willev from tue south seas. 611
Fam. Nannastacidak.
1866. Nannastacidae, Spence Bate, Zoological Record (for 186.")), vol. 2, p. 329.
1878 — 9. Cumellidae, G. 0. Sars, Arrli. Natiirv., vols. S, 4, Middelhavets Cuinaceer,
p. 144.
1880. Cumellidae, Kossmanii, Zoo). Ergebu. Rcise Rothen Meere.s, Malaco.straca,
p. 90.
1887. Cumellidae, Sai"s, Challciigcr llfport.«, vol. 19, Cumacea, p. 62.
18!);>. Nannastacidae, Stebbiiig, History of Crustacea, Iiiteniat. Sci. Ser., vol. 74,
p. 30.-).
189.5. Natinastacidae, Hansen, Isop. Cuiiiac. Stoinat. Plankt(»n-Exp., p. 09.
1900. Nannastacidae, G. O. Sars, Crustacea of Norway, vol. 3, p. 79.
This family agrees with the Canipyla.s)»idae in sevi'ral respects, having no distinct
telson, no pleopods iii the male, inner branch of the uropods one-jointed, the firat and
second peraeopods in the female, but the third and fourth also in the male, furnished
with exopods. In the mouth-organs there are very considerable differences, the man-
dibular molar being blunt in the present family, but acute in the Campylaspidae,
while also in that family the second ma.xillae are formed of a simple plate, but have
the usual subdivisions in the Nannastacidae. In 1896, however. M. Jules Bonnier
described two species of a new genus Procanipylaspis, in which the molar of the
mandible is acute, but the second inaxillac arc iiorniai. Sars in IDOO inclines t(j the
view that Procampylaspis ajiprouchcs nearer to the Nannastacidae than to the other
family. The three pairs of maxillipeds have peculiarities which may relieve the
difficulty by removing Procampylaspis from both the contending families.
Gen. Nannastacus.
1865. Nannastacus. Bate, Ann. Nat. Hist., Ser. 3, vol. 15, p. 86.
1875. Diops, Paulson, Crustacea of the Red Sea (in Ru.ssian), p. 128.
1878 — 9. Nannastacus, Sars, Middelhavets Cumaceer, p. 160.
1880. Nannastacus, Ko.ssmann, Zool. Ergebn. Rothen Meeres, p. 90.
1887. Nannastacus, Sars, Challenger Reports, vol. 19, Cumacea. ]). ()2.
1893. Nannastacus, Stebbiiig, History of C'rnstacea, p. 305.
1895. Nannastncus, Hansen, Isop. Cumac. Stoinat. Plankton-E.\p., p. 59.
This genus i.s distinguished from all other ( 'uniacean geiu'ia at present known by
having two distinct eye.s. At various dates it has had a.ssigned to it the species
uni/uiculatus Bate, longirostris Sal's, Sarsii Ko.ssmann, Sulimli Sai-s, liirsutus Han.son,
and the Diops spinosus and Diops parvulus of Paulson. Ko.ssmann considere it possible
that his own species may be a synonym of Paulson's parvulus. Both sexes have been
described onlj- in the ca.se of unguiculatus, lonffiro.stris, and Sulimii. In reganl to the
fii-st two the .sexual dimorphism so common in the present order is le.ss striking than
usual, but in the third it is strongly accentuated. Hence the (|uestion arises whether
the form described by Sars as the male of his ,V. Suliniii m,iy not really represent
w. V. 81
612 ox CRUSTACEA BROUGHT BY DR WILLEY FROM THE SOUTH SEAS.
a distinct species. This I am strongly disposed to believe to be the fact, since the
two specimens about to be described present parallel differences, but are clearly distinct
species, being both males. Should N. Suhmii prove to be a name covering two species,
it ought to be reserved for the female specimen, since Sars according to his custom
gives precedence to the description of that sex. The male specimens will then require
a fresh designation, in respect of which a suggestion will be made presently.
N.\.N"X.\STACUS OSSIANI n. Sp.
Plate LXIV a.
Carapace tumid, median line sinuous, its concavity flanked by lateral convexities.
Pseudo-rostral projection forming a rather broad, apically rounded process, obliquely
ascending, the distal part concentrically ridged. Antero-lateral corners blunt, very little
produced. Lower and hind margins forming almost a right angle, corner rounded.
Surface reticulate with he.xagonal or .scale-like markings. Fourth and fifth of the free
peraeon segments with dorsal centre strongly raised. Segments of pleon with pellucid
lateral carinae, as if formed by o\erlapping scales, and pair of serrate dorsal carinae
distally produced into a strong tooth in each of the first five segments; fifth segment
not greatly longer than the rest.
Eyes apparently having three facets, which are difficult to distinguish owing to
the darkness of the accompanying pigment.
The first and second antennae scarcely differ from those of jVannastacus iniguicu-
, latii^, described in detail by Sars (Middelhavets Cumaceer, pp. 165, 169), except that
the slender flagellum of the second pair is much shorter, consisting of ten joints
instead of eighteen.
The mouth-organs and peraeopods, as is perhaps sufficiently shown by the figures,
are in close agreement with those of N. unginculatus. The branchial apparatus of the
first maxillipeds was not, however, clearly observed.
The uropods are imperfect, but have peduncles shaped like those of X. unguiculatus,
a little less produced beyond the terminal segment. The spine of the outer ramus is
perhaps relatively a little .shorter, but with the apex broken this remains uncertain.
Length, 2 mm.
Habitat. " Barawon. Auftrieb at night, lO/S/9.5." One specimen, male. Dr Willey
informs me that Barawon, where this plankton capture was made, is a small village at
the entrance to Blanche Bay, New Britain.
Though the shape of this species so much recalls that of iV. Suhmii, that at the
first glance I made little doubt of their identity, this is separated from that not only
by characters of the trunk and pleon, which might be sexual, but also by the pro-
portions of the joints in the fifth peraeopods and outer ramus of the uropods, which
in the one case Sars' figure and in the other his figure and description show to be quite
different.
ox CRUSTACEA BROUGHT 15V DU WII.LEV KlioM THK .SOTTTH SEAS. 613
NaNNASTACUS 0E01!(il li. sp.
Plate LXIV li.
Carapace less tumid than in the ])receding species but with similar depression
and elevations. Pseudo-rostral projection in like manner reminded, but less ob]iijuel\
produced and not ridged on the surface. Antero-lateral corners acute, minutely pro-
duced. Lower and hind margins forming an nbtuse angle, little rounded. Surface as
in preceding species. Fifth peraeon segment and first of pleon with a marked longi-
tudinal medio-dorsal depression. Segments of pleon with pellucid latei-al carinae formed
as in the preceding species, fifth segment considerably the longest.
Eyes larger than in the other species ; antennae similar, but the flagellum of the
second pair much longer, composed of eighteen joints or possibly more, the terminal
joints so excessively slender that their boundaries are diHicult to distinguish. The
mouth-organs agree with tho.se in the companion species. The branchial leaves appear
to bo about .sixteen in number.
The peraeopods are less slender. Especially it will be noticed that in the fifth
pair the antepenultimate joint is little longer than the penultimate, instead of much
longer as in iV. ossiuni.
The uropods differ greatly, for heie tlie peduncle i.s produced beyond the terminal
segment of the pleon at least as much as in iV. unguicidatus, and the outer ramus,
instead of being siibequal in length to the peduncle, as in N. ossiuni, is not half as
long, in eacli case otnitting the terminal spine, which is here of great length. The
inner ramus is very long, slender, spinose.
Length, 2'o mm.
Habitat, the same as that of Nannastacus ossiani.
The specific names are chosen in compliment to Professor Georg Ossian Sars.
Excejit for some differences in the shape of the carapace, N. georgi seems to agree
closely with the male form described by Sars from the Phili])])ines, and if they should
prove to be simply varieties of a single species the name now given will supph' what
is re(iuired for the form described and figured 1)\ Sars.
IS()1'()1).\.
F.\M. Tanaii).\k.
18o3. Taiuiidae (part), Dana, U. S. Expl. E\\>., \ol. 13, pt. 2, p. 71)2.
1857. Asellidae (part), White, Popular Hist. Brit. Crustacea, p. 225.
IStifi. Tananhie (part), Bate and Westwood, Brit. Sess. Crust., vol. 2, p. 117.
1.S.S0. Titnaidiie, Harger, Rep. U. S. Fisheries for 1878, pt. (J, pp. :i04, 41o.
1880. Tanaidue, Sai-s, Lsopoda chelifera. Arch. Naturv., vol. 7, sep. copy p. 20.
188C. Tanaidue, BeddMrd, Challenger Reports, vol. 17, Lsopoda. )i. 11!'.
81—:
614 ON CRUSTACEA BROUGHT BY DR WILLEY FROM THE SOUTH SEAS.
1886. Tanaidae, Sars, Middelhavets Sa.xisopoder, Ai-ch. Xatuiv., vol. 11, p. .30!).
1886. Tanaidae, Norman and Stebbing, Trans. Zool. Soc. London, vol. 12, pt. -i,
pp. 79, 102.
1893. Tanaidae, Stebbing, History of Crustacea, Internat. Sci. Ser., vol. 74, p. 322.
1895. Tanaidae, Hansen, Isop. Cumac. Stomat. Plankton-Exp., p. 50.
1896. Tanaidae, Sars, Crustacea of Norway, vol. 2, p. 10.
1897. Tanaidae. Dollfus, Bull. Soc. Zool. France, vol. 21, p. 207.
1898. Tanaidae, Dollfus, Mem. Soc. Zool.' France, vol. 11, p. 35.
1899. Tanaidae, H. Richardson, Proc. U. S. Mus., vol. 21, p. 819.
1900. Tanaidae, H. Richardson, The American Naturalist, vol. 34, p. 210.
The genus Tanais, on which the name of this family is based, is due to Audouin
and Milne-Edwards who figured a species in 1829.
Leptocheli.\, Dana.
1849. Leptochelia, Dana, Anier. J. Sci., ser. 2, vol. 8, p. 425.
1852. Leptochelia, Dana, Amer. J. Sci., ser. 2, vol. 14, p. 306.
1853. Leptochelia, Dana, U. S. E.xpl. Exp., vol. 13, pt. 2, pp. 792, 800.
1864. Tanais (part), Lilljeborg, Bidrag Sverige och Norrige Isopod. uuderord. och
Tanaid. fam., pp. 7, 11.
1866. Leptochelia, Bate and Westwood, Brit. Sess. Crust., vol. 2 (part 14), p. 132.
1870. Tanais (part), Dohni, Untersuch. liber Bau und Entw. Arthropoden, pt. 2,
p. 141.
1878. Paratanais (part), Harger, Amer. J. Sci., vol. 15, p. 377.
1879. Leptochelia (part), Harger, Pr. U. S. Mus., vol. 2, p. 162.
1880. Leptochelia (part), Harger, Rep. U. S. Comm. Fisheries for 1878, pt. 6,
p. 420.
1880. Paratanais, G. M. Thomson, Ann. Nat. Hist., ser. 5, vol. 6, p. 2, and (1881)
Tr. N. Zealand Inst., vol. 13, p. 207.
1880. Lej^tochelia, Sars, Isopoda chelifera, Ai-ch. Naturv., vol. 7, 1881, sep. copy p. 24.
1881. Leptochelia, Delage, Arch. Zool. exp., vol. 9, pp. 145, 154.
1886. Leptochelia, Sars, Middelhavets Saxisopoder, Arch. Naturv., vol. 11, p. 315.
1886. Leptochelia, Beddard, Challenger Reports, vol. 17, Isopoda, p. 132.
1886. Leptochelia, Norman and Stebbing, Tr. Zool. Soc. London, vol. 12, pt. 4,
p. 108.
1893. Leptochelia, Stebbing, Hist. Crust, Internat. Sci. Ser., vol. 74, pp. 323, 326.
1895. Leptochelia, Hansen, Isop. Cumac. und Stomatop. Plankton-Exp., p. 50.
1896. Dolichochelia, Stebbing, Ann. Nat. Hist., ser. 6, vol, 17, p. 49.
1896. Leptochelia, Stebbing, Ann. Nat. Hist., ser. 6, vol. 17, p. 156.
1898. Leptochelia, Dollfus, Mem. Soc. Zool. France for 1897, vol. 11, p. 40.
1900. Leptochelia, H. Richardson, The American Naturalist, vol. 34, pp. 210, 212,
As already pointed out in 1896, the type species of this genus has the outer
branch of the uropods two-jointed. This character is shared by the .species inimita
ON CRUSTACEA BROUGHT BY DR WILLEY FROM THE SOUTH SEAS. 615
Dana (the type), forresti Stebbing, and lifuensis n. sp. For brasiliensis (Dana) and
Jilum (Stimpson) the inner branch is not described, and the same may be said of
Corsica Dollfus and inermis Dollfus, species instituted in 1898, but for eacii of these
a one-jointed inner branch is to be understood. For inermis it is tiwured beside the
four-jointed inner branch. In Corsica the inner branch is six-jointed, and as M. Dollfus
was half inclined to unite his Corsica either with neapolitana of Sars or with savignyi
(Kroyer) + algicola Harger, in all of which the outer branch has but a single joint, it
may be inferred that the same character belongs to Corsica.
While i-etaining the opinion that Dolichochelia is identical with Dana's genu.s,
I no longer think that Dolichochelia forresti should be united with Leptochelia minuta,
the hand and finger of the chelipeds being very distinct in the two species, which
are also separated in habitat by an immense interval.
Leptochelia minuta, Dana.
1853. Leptochelia minuta, Dana, U. S. Expl. Exp., vol. 13, pt. 2, p. 800, pi. 53,
fisr. o a — d.
1896. Leptochelia minuta (part), Stebbing, Ann. Nat. Hist., ser. 6, vol. 17, p. 158,
From Leptochelia forresti, Stebbing, which in 1896 I thought must be made a
synonym of Dana's species, I now think that L. minuta, is distinct. Upon comparison
of actual specimens some satisfactory marks of difference become available. The
examples in Dr Willey's collection do not agree with Dana's figures either in the
demarcation of the head from the first peraeon segment or in having a short joint
at the base of the first antennae. That was fully to be expected. In other respects
they agree well with Dana's representation, but are distinguished from L. forresti by
the following details. The front of the head is flatly rounded rather than obtuse-
angled, if one may trust mounted specimens for the observation of so minute a detail.
In the first antennae the difference is striking, the second joint in the West Indian
species being four-fifths the length of the first, whereas in L. minuta. it is only
half as long. The third joint, however, which Dana figures as about two-thii-ds the
length of the second, is barely one-fourth of that length or less in the specimens which
I have examined of both species. In the enormously elongate chelipeds L. minuta
has the long fifth joint parallel-sided except at the narrow base, and the elongate
thumb of the sixth joint with a low prominence near the apex, but L. forresti has
a ver}' marked emargiuation near the base of the fifth joint, and has the process near
the apex of the thumb very prominent.
On the number of joints in the flagellum of the first antennae no .stress can
be laid, for one specimen of L. forresti has eight joints and another only six, while
one specimen of L. minuta has six joints on one of these antennae and seven on
the other, but a second has eleven joints in each flagellum. Yet all these
specimens have the remarkably developed chelipeds di.stinguishing the male of this
species.
The two-jointed outer branch of the uropods is a little longer as observed in
616 ox CRUSTACEA BROUGHT BV DR WILLEY FROM THE SOUTH SEAS.
the last mentioned specimen of L. mimita than it is as observed in one of the
specimens of L. forresti, but this may be set to the account of individual variation.
Length, 25 mm.
Habitat. Sandal Bay, Lifu, Loyalty Islands.
It is probable that Paratanais erythraea, Kossmann, 1880 (Zool. Ergebnisse Reise
Rothen Meeres, Malacostraca. p. 103, pi. 7, fig. 1 — 4), is identical with this species.
Leptocheli.\ lifuexsis n. sp. ^
. Plate LXV B.
</. The first two (free) segments of the peraeon are rather shorter than those
which follow. The pleon is a little wider than the peraeon, its last segment bluntly
pointed as usual.
Fhst antennae. First joint twice as long as second, which is twice as long as the
third ; flagellum of six joints, each nearly as long as the third joint of the peduncle, with
perhaps a microscopic seventh joint.
Second antennae shorter than the peduncle of the first, very slender, all the joints
of the peduncle except the last short, the flagellum consisting of one long joint
between two that are quite minute ; the terminal setae long.
Mouth-organs. The upper lip appears to be a narrow plate as shown in the figure.
The oral parts opposed to it showed a group of lobes so small and closely compacted
that I do not venture to discuss their homologies, and in the figure represent what
I saw, perhaps not what I ought to have seen.
First (/natkojMds. These are elongate, but much more substantial than in Lepto-
chelia mimita. The sixth joint or hand can fold back partially into the channelled
margin of the large preceding joint ; its long curved thumb has two strong processes
of the inner margin, separated by a wide interval, and this leaves a large cavity when
the tip of the thumb crosses the tip of the equally long finger; the distal process
of the thumb is lower than the proximal and carries some long setae ; the concave
margin of the finger is seiTate, and armed with ten spinules.
The second gnathopods are of the usual slender form, with the finger nearly as
long as the preceding joint.
The first and second peraeopods resemble nearly the second gnathopods, except
that the finger is much shorter than the preceding joint. The three following pairs
differ by having the second joint stouter, with the sides slightly convex, the fourth
joint not end to end with the fifth, but somewhat under-riding it, the fifth with a
strong spine at the hind apex, accompanied by other smaller spines, and the sixth
with a set of apical spines behind the small curved finger, these in the fifth peraeopods
forming a close fringe of about half a score.
The pleopods appear to have the plumose armature usual in this genus.
The uropods have a peduncle longer than broad, the outer ramus two-jointed.
.'iLLEY. Zoological Results.
prpS
T ii R S D.l
r v,M !»»i . rambndce
A Nannastacus osslan.
B Nannasl:. eorgt.
pto chelld
ox CRUSTACEA BROUGHT BY DR WILLEY FROM THE SOITH SEAS. fi 1 7
scarcely longer than the first joint of the inner, which is five-jointed : the rami are
tipped with long setae.
Length, 2'5 mm.
Habitat. Lifu, Loyalty Islands.
The form shown in Plate LXIV D differs slightly from that above described,
having the processes of the thumb in the first gnatliopods separated by a narrow
interval instead of a wide one, a distinction corresponding to that which Dollfus notes
as separating L. savignyi, Kriiyer, from L. aUjicola, Harger, and L. Corsica, Dollfus,
without making it very sure that all those three .species may not in i-eality be one
and the same.
Habitat. Isle of Pines: from sponge.
Leptochelia lifuensis n. sp. $
Plate LXIVc.
$. In keeping the description of the female separate from that of the male
I am influenced by the circumstance that the specimens assignable to the former
sex attain a size so much greater than is shown by any specimens of the latter, that
doubt as to the identity of the species is not altogether excluded.
Apart from the size, the differences though very considerable are only such as arc
known to occur in the two sexes of this genus.
In the first antennae the stout first joint is three times as long as the second,
the second is but little longer than the narrower third, to which succeeds a minute
apical joint tipped with long setae. Exceptionally in place of the thinl jdint there are
two joints, together not much longer than the single joint.
The .second antennae are much larger than in the male, the fii'st three joints short,
the second and third each tipped at each side with an outstanding pellucid spine, the
fourth joint much longer, having a dark band across the middle ; the short slender
flagellum as in the male consisting of one principal joint between two that are
microscopic, the apical perhaps itself subdivided, tipped with long setae.
The mouth-organs agree closely with the figures and descriptions given by Sars
in 1886 for those of the female of Leptochelia dubiu (Kriiyer). In the first n)axillae
the little crowded apical spines appear to be eleven in number. The backward-directed
palp has an indistinct appearance of being two-jointed, and ends in two nne(iual setae.
The fir.st gnathopods are .stout, the thumb short and thick, with five setules on
its outer margin, the distal part of the inner crenate, the proximal part excavate, the
finger having a prominence of its crenate inner margin corresponding with the
emargination of the thumb, the apex of the finger not (juitc reaching that of the
thumb.
The other limbs agree with those in the male, and the same is true of the
uropods, which have a five-jointed iimer, and a small but distinctly two-jointed outci-
618 ON CRUSTACEA BROUGHT BY DR WILLEY FROM THE SOUTH SEAS.
liwnus. One specimen wa.s observed with four-jointed inner ramus on one side, and a
five-jointed one on the other.
Length reaching .5 mm. or a little over; many specimens much shorter, one with
well-developed marsupium being only 3'.5 mm. long.
Habitat. Isle of Pines : labelled as ' Tanaids from sponges.'
Fam. Anthuridae.
1814. Aiitliiiridae, Leach, Edinb. Encycl., vol. 7, p. 433.
1819. Anthuradae, Leach, in Samouelle's Entomologist's Useful Compendium, p. 107.
1852. Anthurinae (subfam. of Arcturidae), Dana, Amer. Joum. Sci. and Arts, Ser. i,
vol. 14, p. 306.
1864. Anthuridae, Lilljeborg, Bidrag Sverige och Norrige Isopod. underord. och
Tanaid. fem., p. 6.
1866. Anthundae, Bate and Westwood, British Sessile-eyed Crustacea, vol. 2, p. 155.
1880. Anthuridae, Harger, Rep. U. S. Commission Fish and Fisheries, pt. 6 for 1878,
pp. 304, 396.
1882. Anthuridae, Sars, Forh. Selsk. Christian., No. 18, p. 15.
1882. Anthurinae (subfam, of Arcturidae), Haswell, Catal. Australian Malacostraca,
p. 304.
1884. Anthuridae, Haswell, Pr. Linn. Soc. N. S. Wales, vol. 9, pt. 3, and vol. 9,
pt. 4.
1886. Anthuridae, Norman and Stebbing, Trans. Zool. Soc. London, vol. 12, pt. 4,
p. 119.
1886. Anthuridae, Beddard, Challenger Reports, vol. 17, Isopoda, p. 143.
1893. Anthundae, Stebbing, History of Crustacea, Internat. Sci. Ser., vol. 74,
p. 330.
1894. Anthuridae, Chilton, Trans. Liim. Soc. London, vol. 6, pt. 2, p. 209.
1895. Anthuridae, H. J. Hansen, Isup. Cum. u. Stomat. der Plankton-Exp., p. 11.
1897. Anthuridae, Sars, Crustacea of Norway, vol. 2, p. 43.
1900. Anthuridae, H. Richardson, The American Naturalist, vol. 34, p. 215.
In his article " Crustaceology " Leach first of all placed his new genus Anthura
among the Myriapoda (p. 404), but in the supplement to that article he transfers the
family Asellides to the Malacostraca, calling it a tribe, in which the first family is the
Anthuridae (p. 433). For a long time this family did not find acceptance with other
writers, and Lilljeborg in 1864 supposed that he was himself establishing it for the
first time. Milne-Edwards in 1840 combined in a family Idot^ides the genera Arcturus,
Idotea and Anthura, which are now distributed over three families, the Anthuridae
being placed by Sars in the tribe Flabellifera, while the Astacillidae and Idoteidae
stand together in the tribe Valvifera. Besides the authors named in the synonymy
there are two who do not happen to have used the Latin name of the femily, but
who have made important studies for its elucidation, Anton Dohrn in his ' Unter-
suchungen iiber Bau und Entwicklung der Arthropoden,' chapter 5, p. 91, 1870, and
ox CRUSTACEA BROUGHT BY DR WILLEY FROM THE SOl^TH SEAS. 619
Schibdte in the Naturliistorisk Tidsskrift, Sen 3, vol. 10, p. 211, 187.5. Schiodte also
calls attention to the value of Kroyer's description and figures of Anthvra carinata
(Naturh. Tidsskr., Ser. 2, vol. 2, p. 402, and Gaimard's Voy. en Scand., Crust, pi. 27,
fig. .3 a — o). This imperfect but under the circumstances meritorious account was over-
looked by Bate and Westwood. They, like Krriyer, were hampered by want of
material, and were reduced to describe the mouth-organs not of Anthura but of an
essential!}' different genus which they named Paranthiira. The lettering of their figures,
the figures themselves, and the accounts relating to them, involve obscurities, some of
which are acknowledged by the authors, and inconsistencies which it is not so easy
to explain. Schiodte has taken great pains to unravel the tangle, but apparently he
was himself only acquainted with the genus Cyathura. to which Kriiyer's species has
been transferred by Norman and myself In the eleven genera that have been named
within this family, the species have so great a superficial resemblance that agi-ee-
ment in the character of the mouth-organs has no doubt been sometimes taken for
granted, and this the more readily because they are so small and so difficult to dissect.
The paper published in the Transactions of the Zoological Society of London in
1886 was read to the Societ}'' in 1884 and had been in substance prepared several
years earlier. Consequently it omitted from its review of the Anthuridae certain
Australian and New Zealand genera which would otherwise have received notice. The
six genera accepted in that paper are divided between two well-marked sections. In
the first of these, to put the matter briefly, the mandibles and lower lip end obtusely,
in the second acutely. The first section contains the genera Anthura, Cyathura,
Anthelura, Hyssura; the second contains Paranthura and Calathura. The genus Ptilan-
thura, Harger, 1880, is regarded as a synonym of Anthura, but this is an opinion
which I can no longer support.
In Anthura the female has five segments of the pleon coalesced into a single
segment, the mandibles have a three-jointed palp. In Ptilanthura the females have a
distinctly segmented pleon, the mandibles have a one-jointed palp. It is certainly
curious that Harger did not include the latter feature in his generic definition, but
he was a careful writer and had .several specimens at command, so that his definite
statement should scarcely be set aside on conjecture. In regard to the females he
expressly says that they are distinguished from young specimens of Anthura pulita
by the larger eyes and 'the more elongated and distinctly segmented pleon.' An-
thura polita, Stimpson, is in all probability the same as Cyathura carinata (Kroyer),
which agrees with Anthura very nearly as to the coalescence of pleon segments in
the female. Harger named the type species of his genus tenuis on the chance that
it inight prove to be identical with Paranthura tenuis, Sars, 1872, and Sars at one
time accepted the supposed identity, but has now withdrawn his species alike from
Paranthura and Ptilanthura, placing it in a new genus Leptauth ura, which belongs
to the second section of the fiimily, while Harger's genus belongs to the first.
To the eight genera already named must be added Haliophasnia, Haswell, 1880,
and Eisothistos, Haswell, 1884, both from Australia, but there is no information about
the mouth-organs of either to show whether the existing sections of the family are
fitted to receive them. On the other hand the genus Cruregens, Chilton, 1882, from
w. V. 82
(i20 ox CRUSTACEA BROUGHT BY DR WILLEY FROM THE SOUTH SEAS.
New Zealand, was in 1894 made the subject of a very ample discussion by its
author, and this shows that it belongs to the second section by almost all its oral
characters. But there is this to be remarked in the mandibles that, though they
run out to a sharp point distally, as in the other genera of the second section,
this point is here not freely outdrawn, but flanked by 'a thin chitinous plate with
rounded margin,' coiresponding to the serrate blade which flanks the apical tooth or
teeth of the mandibles in the first section. In this genus the mandibles are entirely
without palp, a fact which lends additional probability to the one-jointed palp of
Harger's Ptilanthura. The mandibular palp is indeed very variable in this family,
its third joint being large in Cyathura and Calathura, but very small in Anthura
and Leptanthura. But of all the oral appendages those which give the most trouble
are the second maxillae. For Cyathiira they are clearly figured by Schiodte, in close
connexion with the first pair. For Cruregens Chilton figures them in attachment to
the lower lip. In establishing the genus Calathura Norman and myself say nothing
about them, and recently Sars has expressed the opinion that they are wanting both
in Calathura and Leptanthura. Bonnier figures them for his Calathura affinis, but
that species by the first antennae and by the maxillipeds would rather belong to
Paranthura. In the latter genus Dohm, describing Paranthura nigro-punctata (Lucas),
gives a figure explained as ' coalesced second maxillae (?),' which no doubt, as Dr
Chilton has already suggested, represents the pair- of second maxillae with the deeply
cleft lower lip between them. Now, also, the dissection of a specimen of Calathura
hrachiata, from East Finmark, given me by Canon Norman, shows that they are
present in that species, just as in the species of Paranthura. Should they prove
to be present also in Leptanthura, Bonnier's affinis might find its proper place in
that genus, with which it agrees in the mandibular palp, the uropods, and most
other characters. Bormier, however, does not describe the position of the fifth joint
in the hind peraeopods, which is characteristic in Leptanthura.
From the descriptions available it appears that the maxillipeds are no less variable
than other oral parts, except that the epipod-bearing first joint always seems to be
indistinguishably coalesced with the wall of the head. Omitting this joint from the
calculation, the maxillipeds form only one joint and a rudiment in Cruregens, two
joints in Anthura and Ptilantliura, two and a rudiment in Paranthura, three in
Leptanthura and Cyathura, four in Calathura and Apanthura, five in Anthelura and
Hyssura.
The uropods are a subject of controversy. Chilton and Dohru consider the lower
branch to be one-jointed, not two-jointed, as various authors have stated it to be.
Dr Chilton argues that it would be quite exceptional for the inner branch to be
two-jointed. At the same time he points out that Gerstaecker held the upper branch
to be the inner. If Gerstaecker is right in that interpretation, Dr Chilton's objection
would so far disappear. The question would remain whether in the outer branch we
are to consider that a first joint is consolidated with the peduncle. This seems at
least not improbable.
ON CRUSTACEA BROUGHT BY DR WTLLEY FROM THE SOUTH SEAS. 621
Apanthura, n. g.
PliMiu with segments distinct. Mouth-organs as in Anthura, except maxillipeds,
which have a three-jointed palp, of which the middle joint is much the largest. The
last four pairs of peraeopods, as well as the preceding pairs of peraeon appendages,
have the fifth joint under-riding the sixth.
Name compounded of airo, from, and Anthura, a related genus.
The only genus in the family Anthuridae agreeing with Ajianthura in the structure
of the hind peraeopods is Leptanthura, Sars, from which it is completely distinguished
by the mouth-organs. Anthelura abyssorum, N. and S., 1886, may agree in regard to
the peraeopods with Apanthura, but, if so, it differs from Anthelura elongata, Norman,
and its generic position will require to be reconsidered.
Apanthura sandalensis, n. sp.
Plate LXV a.
Head considerably longer than broad, with a very small projection in the middle
of the front. Seventh segment of peraeon as usual the shortest, not quite so long as
the six following segments of the pleon combined, and these rather shorter than the
oval telson. The body is slightly pubescent, and the fifth pleon segment fianked with
plumose setae.
Eyes small, dark in spirit, near to the front angles of the head.
Upper antennae shorter than the lower, third joint shorter than first, longer than
second, about as long as the three-jointed flagellum, in which the first and third joints
are extremely short.
Lower antennae with the peduncle so commonly occurring in this family, a short
first joint followed by a large somewhat folded second, to which succeed three smaller
joints ; the fifth is here longer than the third or fourth and than the setose three-
jointed flagellum.
Upper lip triangular, broader than long, uns^^mmetrically cleft at the apex.
Mandibles with tridentate apex, but the teeth faintly distinguished ; the pi-ojecting
border between the apex and the feeble molar process is very slightly serrulate ; palp
three-jointed, third joint scarcely so long as first, carrying four or five spines, the
terminal the longest ; second joint with a long spine at apex and a shorter one
below.
Lower lip. The lobes less rounded than in Cyathura and less truncate laterally
than in Anthura, with minvite setules at the apical points and three setae on the
outer margin at the point where it passes from convex to concave.
First maxillae as in Cyathura, the apex bent, its point projecting beyond a row
of closely set spine-like teeth.
Second maxillae closely adpressed to the lower lip, but narrower and shorter, the
outer margin sinuous, the apices apparently bifid, the stems seemingly coalesced.
82—2
622 ON CBUSTACEA BROUGHT BY DR WILLEY FROM THE SOUTH SEAS.
Maxillipeds with the first joint carrying small oval epipo(Js, but not otherwise dis-
tinguishable from the wall of the head, which, as usuallj' in the family, is ventrally
more or less carinate ; second joint not twice as long as broad, seemingly with a
small oval plate near inner angle of apex, which is not produced ; the three-jointed
palp much longer than the basal joint, which is not longer than the second of the
palp; third joint of the palp small, broader than long, attached to the outer part of
the truncate apex of the preceding joint.
First gnathopods. Second and third joints large, broad, third deeply channelled,
fourth joint short, fifth with its triangularly produced point meeting tip of finger, sixth
thick, broad at base, suboval or pyriform, but with setose palm margin almost straight ;
the apex, as in all the limbs, forms a little subcircular expansion beside base of finger;
the finger much curved, narrowing and armed with a seta at about the middle of
the inner margin.
The second gnathopods and first peraeopods are much more slender, the third joint
rather shorter than the second, the fourth short, distally as broad as the length, the
fifth small, triangular, apically almost acute, the sixth narrowing distally, its palmar
margin having a row of setae and an apical spine, with a second row of setae on
the adjacent .surface, the finger about two-thirds as long as the preceding joint, curved,
constricted, with a seta at the constriction.
Second to the fifth peraeopods. These differ little from the two preceding pairs of
limbs, but the fourth joint is rather pyriform than triangular, decidedly longer than
broad, the fifth joint is less narrowly triangular, and the sixth of almost uniform width.
In these limbs the sixth joint and finger show a fine serration of the margins when seen
at a suitable angle.
Uropods. The upper branch is strongly emarginate at the apex, of which the outer
lobe is the naiTOwer; except for two-thirds of the outer margin, this branch is fringed
with setae, some of them very long; it reaches a little beyond the base of the terminal
joint of the lower branch, which is strongly fringed, oval, e.xcept at junction with its
base, which is equal to it in length and breadth and obliquely grooved on the surface.
The telson is oval, about twice as long as broad, slightly carinate longitudinally, with
a few setae on the surface, and several long ones round the apex, which does not reach
the level of the apices of the uropods.
Colour (in spirit) wanting, except for a light brown marbling on the back of head
and each peraeon segment.
Length, 7 mm. Judging from the antennae, the two specimens ai-e probably females.
The one figured has the segments of the peraeon from the third to the seventh
thickened.
Habitat. Sandal Bay, Lifu, Loyalty Islands. Specific name referring to place of
capture.
Gex. Paranthura.
1866. Paranthura, Bate and Westwood, Brit. Sess. Crust., vol. 2, p. 16-3.
1870. Paranthura, Dohrn, Unters. liber Bau und Entwickelung der Arthropoden,
p. 91.
/ViLLEY. Zoological Results
Plate LXV.
TR R5. D«l
Apanthura sand alensis, n. sp.
Leptochelia. lifu ens is . n sp.
ON CRUSTACEA BROUGHT BY DR WJLLEY FROM THK SOUTH SEAS. 623
1886. Paranthu7-a, Norman and Stebbing, Trans. Zool. Soc. London, vol. 12, pt. 4,
p. 122.
1893. Paranthura, Stebbing, History of Crustacea, Internat. Sci. Ser., vol. 74, p. 332.
Several species have been referred to this genus by Haswell and Beddard, but, as
no description is given of the mouth-organs, their generic position remains at present
doubtful.
To the account of the genus given in 188G the following observations maj' be
added. The upper lip, as in Cnlatlmra, is elongate triangular, the pellucid apical part
nearly parallel-sided with rounded tip. The third joint of the mandibular palp is armed
, with a series of spines. The spear-like first maxillae, with reverted teeth on the distal
part, appear generally to protrude in advance of the mandibular apices, as though fitted to
take the lead in piercing a victim. The second maxillae are very delicate organs facing
the lower lip, so as to form a channel, which is greatly strengthened by the mandibles
and the maxillipeds. The only parts which appear to have much freedom of movement
are the first maxillae. When the maxillipeds have been removed, the first maxillae can
be drawn out from between the .second maxillae and the lower lip without disturbing the
other organs. The maxillipeds have a small oval epipod, a long basal (or second) joint,
proximally emarginate at the side for the convenience of the epipod, not, or scarcely at
all, distally produced at the inner margin, being in this respect distinguished from
Calatlmra, as also in the palp, which is tapering, two-jointed, with the division between
the <two joints almost imperceptible, whereas in Calathvra the palp has three joints, the
fii'st very small, the third somewhat lamellar, as shown in Sars' figure of Calathura nur-
vegica. My figure of Calathura bracliiata in 1880 does not give the articulation of the
terminal joint, and shows it edgewise, in its natural position relatively to the rest of the
appendage, but in a misleading one for diagnosis. In Paranthura the maxillipeds clo-sely
resemble those of Leptanthura, but in the new species about to be described the basal
joint is not twice as long as the palp. Also in the new species the distinctness of the
pleon segments medio-dorsally is obscure, though laterally it is clear.
Paranthura lifuensis, n. sp.
Plate LXVI b.
Head a little longer than broad, median rostral point very small, not reaching
so far as the lateral angles. Segments of peraeon not elongate, seventh more than
half as long as sixth, as long as the six following segments of the pleon combined.
Of the latter the first and sixth are distinct, but the intermediate four are ditticult
to distinguish except laterally. Body a little pubescent.
Eyes small, dark in spirit, near to the front angles of the head, the component
ocelli about fifteen in number.
Upper antennae little shorter than lower, third joint longer than second, shorter
than the four-jointed flagellum, in which the third joint is the longest.
624 ON CRUSTACEA BROUGHT BY DR WILLEY FROM THE SOUTH SEAS.
Lower antennae. Second joint rather large, folded; fifth longer than third or
fourth or than the single-jointed flagelluni, which is tapering, fringed with several
tufts of setae.
Upper lij). See remarks on the genus.
Mandibles, as in Calathura, but the third joint of the palp is short, with only
four or five spines.
Lower lip slender and delicate, a longitudinal line down the centre perhaps marking
a cleft, but the length of the cleft, if any, could not be made out. There is a small
tuft of setules at each side of the apex.
First maxillae, very long and slender, with eight minute retroverted teeth along
the distal part. In the dorsal view of the head (fig. C) these organs ^vill be seen
protruding from between the peduncles of the lower antennae, being in fact much in
advance of the mandibles.
Second maxillae. These are slender, narrowing rather abruptly at some distance
from the apex, which carries two or three setules. With the lower lip they form a
narrow pipe. In the figure they are drawn apart to either side of the lip.
Maxillipeds. The epipods are oval, exceedingly small. Not only is the joint to
which they belong indistinguishable, but the following pair of joints is likewise at
its base completely coalesced with its indistinguishable predecessor. The cleft dividing
this pair of unjointed joints is equal to the length of the palp, which, as in Paran-
thura nigroptinctata (Lucas), is curved and tapering, with some setae at the apex,
among which a minute second joint is with difficulty discernible ; a long spine-like
seta tips the inner apical angle of the basal joint.
First gnathopods. Second joint not longer than third, but distally wider and
channelled, fourth much broader than long, cup-like, fifth small, triangular, sixth
broadly pyriform ni general contour, but the palmar margin with a produced point
at its base, between this and the finger having a close-set row of spinules on a
slightly convex region, to which succeed some spines and setae ; the narrow apex
forms a subcircular expansion beside the base of the finger in this and all the limbs
of the peraeon. Finger about two-thirds as long as sixth joint, its concave margin
fringed with a few setules.
Second gnathopods and first peraeopods. Second joint rather longer than third,
fourth cup-like, broader than long, fifth triangular, under-riding the narrowly oval sixth ;
finger as in first gnathopod.
Second to the fifth peraeopods. Second and thii-d joints as in preceding pair,
fourth longer than broad, fifth and. sixth narrow, fifth not under-riding sixth, more
than half as long; finger strongly curved, longer than the fifth joint.
Uropods. Upper branch proximally broad, narrow apex just reaching beyond pe-
duncular portion of the lower branch, in which the terminal joint is not longer than
broad, much shorter than the peduncular part.
Telson elongate oval, wider at the base than at the setose apex, which falls
somewhat short of the uropods.
Colour (in spirit) pallid, faintly speckled on the back.
Length about 8 mm.
Plate LXVf
TR.RS. 0«l
Gnalhia aureola, n sp
EWUim.Ulh Umbridge
Paranthura lifuensis, n sp
ox CRUSTAC'KA BROUGHT liV Dli WILLKV FKO.M TIJE SOl'TH SEAS. 625
Habitat. Saiidal Bay, Lifu, Loyalty Islaml.s. A single specimen. Specific name
referring to place of capture.
Fam. (Inatmiidae.
l«l:i — 14. Gnathonii, Leach, Edmb. Encycl., Ait. Cnustaccolugy, vol. 7, p. SHU
{Gnathianii on p. 402, and referred to as Gnathonii on p. 432).
1814. Gnuthides, Leach, loc. cit., Appendi.x, p. 432.
1S2.J. Deceiiipedes (part), Latn-illc, Fam. Nat. du Regue Animal, p. 289.
1840. Pranisien.s, Milne-Edwards, Hist. Nat. Crust., vol. 3, p. 191.
1847. Pranizidae, White, List of Cru.stacea Brit. Mus., p. 101.
18.")0. Pranizidae, White, Brit. Crust, in Biit. .Mus.. ]i. 73.
18.50. Anceadae, White, loc. cit, p. 74.
18.53. Pranizidae, Dana, U. S. E.xpl. K.xp., vol. 13, p. 791.
18.57. Pranizidae, White, Popular Hist. Brit. Crust., p. 239.
1857. Anceadae, White, loc. cit., p. 243.
1861. Pranizades, P. J. van Beneden, Faune litt. de Belgique, p. 100.
1864. Anceidae, Lilljeborg, Bidrag Sverige och Norrige Isopod. underord. och Tanaid.
fam., p. 7.
1866. Anceidae, Bate and Westwood, Brit. Sess. Crust, vol. 2, pt. 1.5, p. 168.
1880. Gnathiidae, Harger, U. S. Comin. Fish and Fisheries for 1878, i)t. (i, pp. 304.
408.
1885. Anceidae, Sans, Den Norskc Xcjnlhavs E.vp., Crust., vol. 14, ])t. 1, p. .s.5.
1886. Anceidae, Beddard, Challenger Reports, vol. 17, Isopoda, p. 135.
1893. Gnathiidae, Stebbing, History of Crustacea, luternat. Sci. Sen, vol. 74, p. 335.
1897. Gnathiidae, Sai-.s, Crustacea of Norway, vol. 2, pt 3, p. 51.
1900. Gnathiidae, Harriet Richardson, The American Naturalist, vol. 34, p. 214.
Milne-Edwards divided his Pranisiens into two tribes, Pranisiens and Anceeus.
White, who at first included Praniza and Anceus in a single family, subsequently
allotted each genus to a separate fomily. The great distinction belongs to M. Eugene
Hesse of having demonstrated that forms apparently so widely apart, that they could
by good naturalists be placed in separate genera, tribes, or families, were really females
and males of the same species. It is also memorable that Leaoli in 1813 had already
deelare<l his suspicion that this might be the Ciise, while M. Hesse's not readily
accepted announcement of the fact was not made to the Acadtsmie des Sciences in
Paris until November, 1855.
Gen. GN.vniiA, Leach.
1813. Gnathia, Leach, Edinb, Encycl., vol. 7, y. 402.
1816. Ancens, Ri.sso, Hist. Nat. Crust Nice, p. 51.
1818. Praniza, Latreille, Tableau Encyclopediipie et Methodique des trois regnes
de la Nature, pt. 24, Crustacea, pi. 329, f 24, 25.
626 ox CRUSTACEA BROUGHT BY DR WILLEY FROM THE SOUTH SEAS.
1S18. Coelino, Latreille, loc. cit., pi. 336, f. 28.
1825. Anceus, Desmarest, Consid. gen. Cnist., p. 282.
182-5. Praniza, Desmarest, loc. cit., p. 283.
1829. Anceus, Latreille, Regne Animal, nouv. ed., vol. 4, p. 125.
1829. Praniza, Latreille, loc. cit., p. 125.
1840. Praniza, Milne- Edwards, Hist. Nat. Crust., vol. 3, p. 192.
1840. Anceus, Milne-Edwards, loc. cit., p. 196.
1849 Anceus, Lucas, Explor. sci. de I'Algerie, p. 85.
1849. Praniza, Lucas, loc. cit., p. 87.
1855. Anceus, Hesse, Comptes rendus, Novembre 26.
1858. Ancetis, Hesse, Comptes rendus. Mars.
1858. Anceus, Bate, Ann. Nat. Hist., Ser. 3, vol. 2, p. 165.
1858. Praniza, Bate, loc. cit.
1861. Anceus, P. J. van Beneden, Faime litt. de Belgique, p. 102.
1866. Anceus, Bate and Westwood, Brit. Sess. Crust, vol. 2, pt. 15, p. 170.
1870. Praniza {Anceus), Dohru, Untersuch. Bau und Entwicklung der Ai-thropoden,
pt. 1, p. 65.
1874. A7icaeus, Hesse, Ann. Sci. Nat., Ser. 5, vol. 19, Art. 8, p. 8.
1880. Gnathin, Harger, U. S. Comm. Fish and Fisheries, pt. 6, p. 410.
1880. Anceus, Kossmann, Raise in die Klistengebiete des Rothen Meeres, Malacostraca,
p. 105.
1884. Anceus, Hesse, Ann. Sci. Nat, Ser. 6, vol. 17, Art. 6, p. 1.
1885. Anceus, Sars, Den Norske Nordhavs Exp., Crust., vol. 14, pt. 1, p. 85.
1886. Anceus, Beddard, Challenger Reports, vol. 17, Isopoda, p. 135.
1893. Gnathia, Stebbing, History of Crustacea, p. 337.
1897. Gnathia, Sars, Crustacea of Norway, vol. 2, pt. 3, p. 51.
Various other references to the voluminous literature vnW be found in Bate and
Westwood's work. Leach in 1813 placed the family which he instituted for this
single genus between the ' Squillarii ' and the ' Gammarini.'
It was this probably that induced Latreille in 1829 to neglect the guidance of
Desmarest and to place the genus, not near the Amphipoda, but actually among them.
Risso in 1816 had taken the still more remarkable step of placing it in his family
' Paguriens ' between Hippa and Parjurus, and was rewarded for his absurdit}' by a
prolonged upholding of his name Anceus over the earlier given Gnathia. Upon Leach
has been saddled the synonym Praniza, which he not only never published but prob-
ably never wished to publi.sh. It was given to the world in a haphazard manner
by Latreille in the great Atlas to the Encyclopedic Methodique, in which Slabber's
figure of Oniscus marinus is copied and stated to belong to Dr Leach's genus Praniza.
For the date of 1769 at which Slabber's original figure appears to have been published,
it must be considered a highly creditable production, and that it belongs to what is
often called the Praniza-iovro. of the genus Gnathia is beyond question.
In this genus the absence of the fifth peraeopods is not limited to the young,
but extends to the adults of both sexes.
UN CRUSTACEA BKUUGHT BY DU WILLEV FROM THE SOUTH SEAS. 627
Gnathia aureola, n. sp.
Plates LXVIa and LXXIV e.
The inconvenience must be admitted of publishing a new species, of whicli the
fully developed form is not known in either sex. But in the present instance Dr
Wiiley took special pains to preserve an exact record of the colouring in life, and, as
this is rather remarkable, it would be a pity to leave it associated with an unnamed
larva. For future use it may be well to recall the peoiliarity on which Hesse several
times emphatically insists, that syjccies of Gnathia parasitic on fish do not tvssume the
sexual forms while on tiie fish, but that, when removed from their host, if kept in
sea-water, they sooner oi' later do assume these forms. Hesse's own experience was
that the larger or full-grown larvae moulted almost immediately after removal. At
the same time he warns the experimenter that these little captives have a surprising
agility and are sui'e to escape unless special precautions are taken.
In general appearance the present form differs little from the corresponding stage
of Gnathia maxillans, the small, delicate peraeopods forming an absurd contrast to the
massive fifth and sixth segments of the peraeon which offer considerable resistance to
the impact of a penknife and cut like a piece of hard cheese.
The first antennae have the third joint considerably longer than the fiist and
second combined, those two being together about as long as the slender flagellum,
in which the second joint is longer than the first plus the third and fourth, the
second to the fourth carrpng sensory filaments. In the second antennae the last juint
of the peduncle is as long as the two preceding combined, but a little shorter than
the slender 7-jointed flagellum.
Over the mandibles lies a broadly triangular piece, distall}' deeply emarginate,
with a narrowly oval central process between the rounded corners of the emargination.
This I sui)pi)se to represent the epi.stome and upper lip.
The mandibles have about nine microscopic teeth on the incurved narrow apex,
the iiarniw portion being longer than the broader proximal part, while in the mandibles
f)f Anceus maaillaris the reverse is the case.
The first maxillae are extremely narrow, except quite at the base, with three tiny
teeth near the needle-pointed apex. The second maxillae are similar, but rather shorter
and not quite so slender.
The maxillipeds have a tolerably even breadth till near the aj)ex, the distal lobe
being beset with setules and apically carrying two little teeth or spinules.
The first gnathopods though leg-like ha\e niurh the eharaclir of mouth-organs.
Thi'ir hooked nails project in front on either side of the organs abovi' described. The
]H iiultiuiatc joint is distinguished from the small triangular antepenultiniate, which
undii-rides it, by a faintly perceptible suture. The preceding joints are rather short,
sub('i|ual. Tile five following pairs of limbs are all very similar one to anothor, the
last rather the longest; the armature is very slight, its most conspicuous features
being a subapical spinule on the bul>;ing margin of the third joint, spinules at apices
of thi' thvvr following joints, which also have the straight margin microscopiailly serrate,
W. V. 83
628 ON CRUSTACEA BROUGHT BY DR WILLEY FROM THE SOUTH SEAS.
with a spiuule at about the middle of this serrulation in the penultimate joint. The
finger has a setule adjoining the nail on the concave margin.
The pleopods have peduncles broader than long, their inner margins approximate,
connected by two pairs of coupling spiues. The outer ramus is about as long as the
peduncle, little longer than broad, fringed distally with nine plumose setae. The inner
ramus is rather longer, not broader, fringed with eight setae.
The uropods have the rami subequal, not quite twice as long as broad, the inner
with si.x, the outer with four, plumose setae. The outer has also three setules at the
outer corner.
The telson is triangular, the length equal to the breadth at the base, with a
setule near middle of each lateral margin, and an apical pair.
Numerous specimens were obtained by Dr Willey at Lifu, Sandal Bay (near Kiki),
on the 17th of February, 1897, parasitic on gills of the white ocellated 4-spined sting-
ray, Aetiobatis narinari, attached both to the gills and to the walls of the gill-
chambers. There were many of the larger and a few of the smaller specimens. Dr
Willey says, " The swollen portion of body (mesosoma) of former was light gold and
black — gold prevailing in living condition — characterised also by two large lateral
golden orioles on each side and one anteriorly on each side of front and of mesosoma.
The mesosoma darkened very much in alcohol, and the gold rings and spots faded to
a pale gi-eenish tint. Abdomen yellowish white." He adds that the eyes were dotted
with gold spots, that a black longitudinal line lay immediately below the points of
insertion of the limbs on the mesosoma, that the whole of the dorsum was covered
with gold spots, usually aggregated into area-like groups, but leaving free parts of a
medio-dorsal black band. The large hinder gold rings included gold spots with a
central black one.
The total length is or, mm., length of thickened part of mesosoma or peraeon
3'5 mm., and its height 2 mm. The few small specimens, though about three-quarters
as long as the large ones, were very much below them in total bulk.
The specific name refers to the golden circlets of the living colour. Slight ti-aces
of these remain only in the small specimens.
The seventh peraeon segment is not in this species clear of the sixth as in
Anceus Rhinubatis, Kossmann, from the Red Sea. Kossmann speaks of the second
antennae as the front, and of the first as the hinder.
Fam. Cirolanidae.
1880. Cirolanidae, Harger, Rep. U. S. Comm. Fisheries for 1878, pt. 6, pp. 304,
376.
1890. Cirolanidae, H. J. Hansen, Vid. Selsk. Skr., Ser. 6, v. 3, pp. 27.5, 310,
317, 318.
1893. Cirolanidae, Stebbing, History of Crustacea, Intemat. Sci. Ser., vol. 74,
pp. 341, 342.
189-5. Cirolaiiinae, Hansen, Isop. Cumac. Stomat. Plankton-Exp., p. 12.
1897. Cirolanidae, Sars, Crustacea of Norway, vol. 2, p. 68.
ON CRUSTACEA BROUGHT BY DR WILLEY FROM THE SOUTH SEAS. 629
1899. Cirolanidae, Hairiet Richardson, Proc. U. S. Mus., vol. 21, p. 822.
1900. Cirolanidae, Harriet Richardson, The Ameiican Naturalist, vol. 34, p. 215.
Hansen in 1890 included in this family Leach's three genera Cirolana, Gonilera
and Eurydice, together with Batltynomus A. Milne-Edwards, and Anuropus Beddard,
for the latter of which in 1893 I suggested a separate family Anuropidae. To the
Cirolanidae has since been added the genus Cirolanides, Benedict, from ft-esh water,
and I am now proposing a new genus Hansenolana.
Gen. Cirolana, Leach.
1818. Cirolana, Leach, Diet. Sci Nat., vol. 12, p. 347.
1840. Cirolana, Milne -Ed wards, Hist. Nat. Crust., vol. 3, p. 235.
1867. Cirolana, Bate and Westwood, Brit. Sess. Crust., vol. 2, p. 294.
1890. Cirolana, Hansen, Vid. Selsk. Skr., Ser. 6, v. 3, pp. 318, etc.
1893. Cirolana, Stebbing, History of Crustacea, Internat. Sci. Sen, vol. 74, p. 342.
1897. Cirolana, Sars, Crustacea of Norway, vol. 2, p. 69.
1899. Cirolana, H. Richardson, Proc. U. S. Mus., vol. 21, p. 822.
1900. Cirolana, H. Richardson, The American Naturalist, vol. 34, p. 215.
Many additional references will be found in the works above cited.
Cirolana pleonastica, n. sp.
Plate LXVII A.
This species is most nearly allied to Hansen's Cirolana sulcata from the Cape
of Good Hope, but well distinguished from that species by the different sculpturing
of the pleon, and by the setose surface of the outer ramus of the uropods.
First segment of the peraeon considerably the longest ; all the peraeon segments
marbled with brown spots, which above the hind margin form a transverse row,
but with a clear central space running lengthwise ; last four pairs of side-plates
strongly sulcate, last two a little produced, subacute. First segment of pleon concealed;
sides of fourth with obtuse corners completely overlapping but not concealing those
of the fifth. Terminal caudal plate triangular, from the middle fringed with plumose
setae, eight spines encircling the rounded or almost truncate apex ; down the centre,
producing a sulcate appearance, are four to five pairs of processes successively smaller;
a dark centre gives the process the look of a tooth, but on nearer inspection it is
found to be ovate, projected backward. Similar processes are found, twelve in number
but minute, above the hind margin of the last peraeon segment, and in a similar
position but not quite so small, to the number of nine on the fourth, and of seven
on the fifth pemeon segment. In profile the hinder portion of the animal has a
somewhat serrate outline.
83—2
630 ON CRUSTACEA BROUGHT BY DR WILLEY FROM THE SOUTH SEAS.
Eyes rounded, with thirty to forty ocelli. In one set of specimens dark brown,
in another set black.
First antennae scarcely as long as peduncle of second, tliird joint a little longer
than first or second, the latter two faintly separated; flagellum equal in length to the
peduncle, ten-jointed, with hyaline filaments. In the specimen dissected the first joint
of the flagellum was extremely short on one antenna, on the other it was the longest
of all ; each flagellum was ten-jointed.
Second antennae nearly reaching the end of the fourth peraeon segment, fourth
and fifth joints of the peduncle subequal, flagellum nearly twice as long as the
peduncle, twenty-seven-jointed, sparingly ciliated.
Upper lip short but broad, rather strongly emarginate.
Mandibles. The left mandible shows on the cutting edge four distinct teeth,
though the two in the centre have their broad edges almost in a continuous line ;
in both mandibles the plate between the cutting edge and the molar has two or three
of the spines rather conspicuous; the third joint of the palp is shorter than the first,
broad, except at the truncate apex.
First maxillae. The inner plate has the usual three thick plumose setae, of
which the lowest is the longest; the outer plate has twelve spines, all slender, three
or four very slight, one or perhaps more a little dentate, and on the plate's inner margin
are four minute teeth or spinules. As usual, the muscles of these appendages are very
powerful.
Second maxillae. The inner plate has the broad oblique distal margin fringed
with slender setiform spines and three plumose setae below, of which the lowest is
the longest ; the other two plates, which in Hansen's later view belong in common
to the third joint of the maxilla, are naiTOw, rather long, carrying several apical
setae, the inner having setae also on its inner margin.
Maxillipeds. These have the fifth joint much wider than the fourth, with seven
setae on its outer margin, and its inner apex a little emarginate ; the sixth and
seventh joints each have five setae on the outer mai-gin.
First gnathopods. The fourth joint on its inner margin has six short blunt-
headed spines attached to the inner surface, four ordinary spines attached to the
outer surface: the fifth joint is triangular, small, almost embedded in the inner
surface of the fourth, and somewhat under-riding the sixth, which has on its inner
margin three spines, the apical the largest.
Second gnathopods. These are rather longer and more slender than the first;
the third joint has at the inner apex two spines, the upper of which is button-like,
scarcely at all projecting; the fourth joint has four blunt spines, and a little apart
from them a stout apical spine ; the fifth joint is short, not overlapped by the fourth,
and not under-riding the sixth.
First peraeopods. These closely resemble the second gnathopods.
Second to fifth peraeopods. These are nearly alike except in length, the fourth
being the longest; all having numerous apical spines on the thu-d, fourth, and fifth
joints; the second joint is smooth except in the fifth pair, which has a scanty supply
of setae on its outer margin.
WlULEY. ZoOI-v.
ON CRUSTACEA BROUGHT BV DR WILLEY FROM TllK SOUTH SEAS. 631
Pleopods. The peduncle is broader than long, with four uncinate spines on the
inner margin of the first and second pairs, 6 in the third, 3 in the fourth, none in
the fifth ; below the hooks in the second pair there are five spine-like setae. In the
fii-st four pairs both rami are partially fringed with plumose setae ; in the last three
pairs the outer ramus has a transverse suture. The male appendage of the second
pail' is straight except at the point of attachment, and has a subacute apex, which
reaches little beyond the inner ramus. In the fiftli pair the peduncle has a pointed
outer apex, and the inner ramus sends up a process in front of the jjcduncle's inner
margin.
Uropods. The peduncle strongly produced, its apical part fringed on the inner
margin with plumose setae ; the inner ramus reaching beyond the telson, its broadly
rounded, strongly fringed distal part having eight or nine spines among the plumose
setae ; the outer ramus much narrower and slightly shorter ; its outer margin coarsely
serrate, nearly straight, with a few spines and setae, the apex a little notched ; the
inner margin and the distal surfaces crowded with plumose setae.
Length, 8"5 mm. ; breadth rather more than a third of the length.
Habitat. New Britain, Blanche Bay, at 100 fathoms depth, and at 60 fathoms.
Those from the greater depth, as preserved, have brown eyes and dark dorsal markings,
those from the smaller depth have black eyes, but are otherwise quite pale in colour.
The specific name refers to the richness of detail in the pleon, beautiful to observe,
but rather too much of a good thing for the efforts of an ordinary pencil.
ClKOLANA ALBICAUUATA, U. sp.
Plate LXVIIb.
Head broader than long, little immersed, rostral point minute. Segments of peraeon
broad, not very unequal in length, first the longest. Anterior side-plates oblong, those
of seventh segment produced acutely backward, but not beyond the first segment of
the pleon, which is well displayed. Second to fourth segments of pleon slightly, and
fifth considerably, narrower than first. Terminal segment below insertion of the uropods
narrowing with gently convex sides to a subacute apex, its lower part fringed with
long plumose setae and eight spines.
Eyes large, dark, distant, subquadrangular, broader behind than in front, ocelli more
than fifty in number.
First anteimae short, moderately stout, second joint narrower than first or third,
flagellum shorter than peduncle, tapering, seven-jointed.
Second antennae about half as long as the body, peduncle (as in Girolana gracilis,
Hansen) having the third joint wide.st distally and the fourth widest proximally, the
two subequal, each shorter than the fifth; fiagellum twice as long as peduncle, with
about twenty joints, many of them slender.
The left mandible has the two middle teeth of its cutting plate very fiat and
feebly separated. All the joints of the mandibular palp aie narrow. The firet maxillae
have the usual three stout plumo.se setae on the inn<r ])late; the outer is apically
fringed with ten spines of unequal length, all slender, some pectinate. The maxilli|)o(ls
632 ox CRUSTACEA BROUGHT BY DR WILLEY FROM THE SOUTH SEAS.
have the fifth joint rather broader than long, with four or five setae on its outer
margin.
First gnathopods. The third joint is tipped with long setae on the outer apex,
the fourth has its outer apex narrowl}- produced along the base of the si.xth and
tipped with slender spines or spine and spiniform setae, its inner margin canying three
stout spines and others of slighter dimensions ; the fifth joint is quite small, under-
riding the si.xth, which has a row of four spines along the inner margin : the finger
is long, tipped with a short nail.
The second gnathopods differ from the first in that the fifth joint is rather larger,
not under-riding the si.xth, and the outer process of the fourth only reaches the end
of the fifth : the sixth joint has three spines along the inner margin. The finger is
as long as the fifth joint.
First peraeopods in close agreement with the second gnathopods.
The second to the fifth peraeopods agree in general structure, but \vith consider-
able differences iii size and other details. The second are much the smallest, and
agree with the third in having the second joint narrowly oval, with very fine marginal
and apical setae, while in the fourth and fifth pairs this joint is broad, especially at
the lower part in the fifth pair, and is fringed with plumose setae all along the hind
margin and with very long ones on the front apical margin. In all the pairs the
third, fourth, and fifth joints are strongly spined ; these joints are successively longer
to the fourth pair, but in the fifth scarcely so long as in the third ; the sixth joint
is longest in the third pair and shortest in the fifth.
The second j^leopods in the male have the stiliform process obtuse at the apex,
and not quite so long as the inner ramus. The rami of the fifth pair are very broad ;
in the outer the transverse suture is only faintly perceptible.
Uropods. The peduncle, with a few setae on its inner margin, is produced on the
inner side well to the middle of the inner ramus, which is more than twice as broad as
the outer, and without being much longer reaches much beyond it, as well as somewhat
beyond the telson. On the distal half or two-thirds it is fringed with long plumose setae,
intermingled with spines, five on the convex serrate inner margin, and three on the much
straighter outer margin, these margins meeting in a subacute apex. The slender outer
ramus is similarly armed.
' The whole of the back is thickly sprinkled with dark stellate markings to the end
of the fifth pleon segment, the compound terminal segment except just at the base and
the uropods being quite clear of markings and colour, a peculiarity to which the specific
name alludes.
Length, 4'.5 ami., breadth nearly half the length.
Habitat. Barawon, Blanche Bay, New Britain.
The species that most nearly approach the present one are Cirolana neglecta,
Hansen, from the Mediterranean, Cirolana gracilis, Hansen, probably from the West
Indies, and Cirolana latistylis, Dana, from Straits of Balabac, north of Borneo. C. neg-
lecta is thrice as long ; C. gracilis, which is 8 mm. long, has the second joint of the fifth
peraeopods no wider than that of the thii-d ; so that these cannot well be confused with
the species above described. Dana's species, however, being only ' three lines long ' or a
ON CRUSTACEA BROLKiHT BY DR WILLEY FROM THE SOUTH SEAS. 633
quarter of an inch, is not put out of court by any great superiority of size. Unluckily
Dana has only given a rather meagre description of it and no complete figure. But he
states that the first pleon segment is nearly concealed under the peraeon, that the uropods
do not reach beyond the telson, and that their setae are not half as long as the rami. In
the.se respects, therefore, it differs from our species. His account of the legs is obscured
by what must be a misprint. He says of them that ' the fourth joint of the third
pair is a little shorter than either the third or fifth pairs, and longer than the tarsus.'
If the word 'pairs' has slipped in by mistake where 'joint' was intended, the passage
would mean, according to our enumeration of limbs and joints, that in the first
peraeopods the fifth joint is shorter than the fourth or the sixth, which would be a
character common to several species, but it would further mean that the fifth joint
is longer than the seventh. This interpretation will agree with Dana's figure of the
limb, but that figure shows the fourth joint only a little way produced over the outer
margin of the fifth, instead of to its extremit}' as in our species.
ClROL.iNA ORIENTALIS, Daua.
1853. Cirolana orientalis, Dana, U. S. Expl. Exp., vol. 13, pt. 2, p. 773, pi. ol,
fig. 7 a — d.
1890. Cirolana orientalis, Hansen, Cirolanidae, p. 117, pi. 4, fig. 1 — ih, in
K. D. Vid. Selsk. Skr., Ser. 6, vol. 3, p. 353.
As pointed out by Hansen, there are two striking features in this species.
" The head is produced in front into a large, advanced process, apically dilated,
coalesced with the frontal plate, and truncate in front," and the uropods are peculiar,
the inner ramus being emarginate on its outer edge, and the outer ramus, which is
the longer, having its outer edge bai'e. Dana's figures show the cephalic process but
ignore the emargination of the uropod, which was perhaps regarded as an accidental
malformation of the ramus. The terminal segment has a pair of well-marked pits
or depressions near the base.
Hansen describing a subadult female, gives fifteen joints for the flagellum of the
first, and twenty-one for that of the second antennae. His specimen was 10\S mm.
long. In a specimen 12 mm. long, I find the flagellum of the first antennae having
on one side seventeen, on the other only twelve joints, in each case the joint next the
peduncle being extremely short. Corresponding to these two respectively, the flagella
of the second antennae had twenty-four and twenty-five joints.
The cutting edge of the right mandible is formed of three large approximate
teeth, the uppermost rounded, the other two triangular. In the left mandible between
the rounded upper and the triangular lower tooth there is a broad low tooth giving
a very different appearance to the cutting edge of this organ.
In the maxillipeds the second joint is much the longest, apart from its produced
plate, which is also long, armed with several plumose setae and near its rounded
apex with a single strong hook ; the fifth joint is much larger than the sixth.
The specimens were labelled as surf isopods, Conflict Group, New Guinea. A single
specimen was from ' Isle of Pines.'
634 ON CRUSTACEA BROUGHT BY DR WILLEY FROM THE SOUTH SEAS.
CiROLANA MINUTA, H. J. Hansen.
1890. Cirolana minuta, Hansen, Cirolanidae, p. Ill (K. D. Vid. Selsk. Skr. p. 347),
pi. 3, fig. 5—5 d, pi. 4, fig. 1—1/.
The frontal plate has an acuminate horn at its base, a distinctive character
but not especially easy to observe. Hansen describes and figures the peduncle of
the first antennae as two-jointed. In the specimen here referred to his species
there are two short joints followed by a long one, the first joint the thickest ; the
flagellum has nine joints, the last two very small. In the second antennae one of
the flagella has seventeen joints, the other twenty-one ; Hansen gives seventeen or
eighteen joints.
The right mandible has the ujjper tooth of the cutting edge blunt, the next
triangular, short, the lowest triangular, long; in the left mandible the blunt upper
and sharp lower tooth are both small, with a rather long low obscurely bipartite
ridge between them. The other mouth-organs are in close agreement \vitli Hansen's
figures. The vibrating plate of the second joint of the maxillipeds has only about
eight setae on the outer margin, and they are rather coarsely plumose. The large
size of the fifth joint is characteristic.
In this species the second joint of the sixth and seventh paraeopods is not
adorned with long plumose setae.
In the uropods the inner ramus is longer and much broader than the outer ;
both have the apex bifid. Under slight pressure the inner ramus reaches very
distinctly beyond the last caudal segment, whereas Hansen speaks of that segment
as reaching a very little beyond the uropods, no doubt indicating the appearance without
pressure.
Length of specimen, 3"75 nan.
Habitat. Sandal Bay, Lifu, Loyalty Islands.
The possibilit}' is open that this species may be identical with the incompletel}'
described Cirolana latistylis, Dana.
, Hansenolaxa, n. g.
Mouth-organs as in Cirolana. Head wider in front than behind. Segments of
pleon widening to the fourth, which conceals the angles of the fifth. First gnathopods
complexly subchelate, much larger than an}- of the following limbs of the peraeon.
All the pleopods with the peduncle broader than long and with both rami mem-
branaceous, the peduncle of the first pair with eight uncinate spines on the inner
mai'gin.
As well from Cirolana as fi'om its neighbouring genera Conilera and Euri/dice
the new genus is distinguished in a marked manner by the character of the first
limbs of the peraeon. Among the species of Cirolana the aberrant C. sphaeromi-
formis, Hansen, makes the nearest approach to the new genus by the general shape
and some peculiarities of the head and pleon.
ON CRUSTACEA BROUGHT BY DR WILLEY FROM THE SOUTH SEAS. 635
The name of the genus is framed in compliment to H. J. Hansen, whose work
has shed so much light on the structure of the Cirolanidae and the nearly related
families.
HaNSENOLANA AXISOPOUS, n. sp.
Plate LXVIII a.
Head twice as broad as long, more than half as broad as the peraeoii, deeply
immersed, front broader than the straight hind margin, truncate, but with small rostral
point and the lateral angles rounded, surface wrinkled. Peraeon only slightly convex,
first segment the largest, wrinkled, its front angles much produced, rounded, second
segment the shortest, the rest subequal, with hind margins faintly nodulose, the side-
plates successively more acute and reaching further beyond their respective segments,
those of the seventh .segment overlapping the fourth pleon segment completely. Pleon
broad, widening to the fourth segment, first segment and angles of second and fifth
concealed, angles of the fourth reaching the apices of the side-plates of the seventh
peraeon segment, fifth segment at the middle rather longer than the fourth. Terminal
segment with a breadth at its base more than twice its length, which exceeds that
of the four preceding segments combined, sides curved, apex slightly curved, less than
half the basal breadth. Earlier segments of ploon have a median lobe which narrows
to the base of the terminal segment, thence to the apex forming a low carina. By
help of the second antennae above and the uropods below the whole body is strikingly
parallel-sided ; many parts of it carrying small hairs.
E\'es placed at the antero-lateral corners of the head, small, but with several
(about eighteen) ocelli.
First antennae shorter than the peduncle of the second, very slender, first joint
the longest, width not uniform, second shorter and much narrower, followed by what
appears to be an extremely short third peduncular joint, flagellum seven-jointed, a
little shorter than the first, a little longer than the second, joint of the peduncle. But
this account must be modified if, as is quite probable, the first joint of the peduncle
is composite, representing the first and second joints in coalescence.
Second antennae. Peduncle much stouter than in first pair, third and fourth joints
equal, fifth a little longer than either, flagellum slender, as long as the jjeduncle,
twenty-one-jointed.
Frontal lamina widening to the rounded apex, its base perhaps concealed by a
rounded median projection of the wide, very short epistome. Upper lip broad and short.
Mandibles. The trunk from a broad base narrows to the cutting edge, which on
the right mandible is divided into three strong tinequal teeth, on the left has a
couple of denticles at the top and a tooth below. //; situ, as described by Hansen
for the Cirolanidae in general, the cutting edge of the right mandible is covered by
that of the left. The blade-like molar has the usual row of teeth within the thin
convex front edge ; between this and the cutting edge is the part called ' lacinia
mobilis' by Hansen, here sun'ounded by eighteen spines and preceded by a thin, slightly
spinulose lobe, apparently attached to the cutting edge. Second joint of palp longer
w. V. 84
(536 ON CRUSTACEA BROUGHT BY DR WILLEY FROM THE SOUTH SEAS.
than first or third, not strongly armed, third joint with an even row of short spines
and one or two long ones at the apex. Anterior articular condyle not very prominent.
First maxillae. Inner plate with three apical setae or spines and a slender sub-
apical one ; outer plate with nine strong spines distinguished.
Second maxillae. Inner plate (lacinia of second joint) apically fringed with about
eight spinules and two long feathered setae ; outer plates (lacinia of third joint divided
into two branches, according to Hansen) are very short, each surmounted with a
couple of setae : of these plates, so far as I can discern, only the inner is here
articulated.
Maxillipeds. First joint forming a small setose projection in front of the second,
its epipod forming a broad projection behind it, this also carrying four small setae ;
second joint a little longer than broad, its plate carr3ring one or two hooked spines
on the inner margin, a simple spine on the surface and plumose setae at the apex ;
thu'd joint broader than long, as also arc the three following joints, of which the
first is cup-shaped, the second the largest, and all have marginal setae; the seventh
joint is narrowly oval, with setae on the hind margin and apex.
First gnathopods. These, when in situ, largely conceal the mouth-organs, the third
joints meeting just over the peduncular part of the maxillipeds. They are much
more massive than any of the following limbs, which on the ventral surface of the
animal are wide apart. Second joint substantial, more than twice as long as broad,
third about half as long, fourth short but wide, receiving in a sort of cup the
somewhat cup-shaped short and wide fifth juint, which on the inner margin forms a
blunt tooth or lip, over which the long and strong finger projects, the broad oval
sixth joint having its inner margin broken into two tooth-like processes, each like
the lip of the fifth joint carrying a minute spine. The arrangement of these last
three joints produces a ' complexly subchelate ' gi-asping apparatus, in which the blunt
inner apex of the fourth joint may perhaps take a share, for the point of the finger
appears to approach it near enough to assist in holding an intervening object.
Geaerally in the Cirolanidae the first three pairs of limbs are in near agreement.
Second gnathopods, with all the joints attached end to end, none of them notably
setose or expanded or under-riding or over-riding a succeeding joint. Second joint more
than twice as long as thii'd, which is subequal to fourth plus fifth, fourth longer
than fifth, with three short stout spines near the base and three near the apex of
its inner margin, fifth joint with one such spine at the inner apex, sixth joint longer
than fifth or than the little curved bidentate finger. In Girolana horealis (Lilljeborg)
the fourth joint over-rides the fifth, in C. sphaeromiformis the fifth under-rides the sixth.
Fifth peraeopods. These differ little from the second gnathopods, except that the
second joint is rather shorter and stouter, the fifth joint longer, the sixth narrower
and not longer than the fifth. The fourth and fifth joints have one or two spinules
at the apex on each margin, but the fourth joint is without the stout spines ob-
servable on the inner margin of the second gnathopod.
Pleopods. The peduncle of the first pair carries eight uncinate spines, of the second
six or seven with two slender spines or setae, of the third five, of the fourth three
uncinate spines with two plumose setae, the spines being longer than those of the
WlLL,BY. ZoOLOaiC>
Plate LXVDL
TRRS nti
Hansenolana anisopous. n, g. et sp
"TUy)
E °<VUsan .CaubncLge
Renocila periophthalmi. n sp
ON CRUSTACEA BROUGHT liV DK WIM.in I'ROM THE SOUTH SEAS. 637
preceding pleopods. The peduncles of the fifth pair are less prominent than those
of the fourth and are perhaps unarmed. The plates of the first and second pleopods
are uanowcr than those of the following pairs, and both inner and outer plates have
some plumose setae. In all the inner plate is smaller than the outer; in the last
three pairs it is without setae, and in these pairs the outer plate has a transverse
suture.
Uropods. The peduncle has its inner margin acutely produced to about the
middle of the ver}' broad inner ramus, which does not reach the end of the pleon.
The outer ramus is rather more tlian twice as long as broad, oblong, with rounded
apex, shorter than the inner ramus, which widens distally till it is twice as wide
as the outer. Both curve a little inward.
Length, 10 ram., breadth, 5 mm.
Habitat. Isle of Pines, south of New Caledonia.
The specific name, from the Greek liviTOTrov^, uneijual-footed, refers to the charac-
teristic size of the first legs, which in this family is rather remarkable. The spelling
anisopous is preferred to anisop2is, to precludi' if possible the atrocity of anisopa,, as
a supposed correction to agree with the generic termination.
Fam. Ai.cirdxidae.
1890. Aldronidae, H. J. Hansen, Vid. Selsk. Skr., Ser. C, vol. 3, pp. 285, 312, 390.
1893. Aldronidae, Stubbing, History of Crustacea, Internal. Sci. Ser., vol. 7-1,
pp. 341, 34f!.
To this family Hansen assigns his own two genera Alcironu and Lanocira, Tacliaea
of SchiiJdte and Meinert, and possibly Kossmann's Gonlana. The limbs of the peraeon
are withotit long natatory setae.
Gen. Ai.cikona, Hansen.
1890. Alcironu, Hansen, loc. cit., pp. 313, 391.
1893. Alcirana, Stebbing, loc. cit., p. 346.
The cl^^eus is broadly crescent-shaped. The genus contains apparently four species,
krehsii and insularis of Hansen, together with that named .Ega multidigitu by Dana,
and that named Cirolana multidiqitdtc by Miers.
Ai.ciKo.NA iNSLLAius, Hansen.
1890. Alcironu insularis, Hansen, Cirolanidae, pp. .51, 1.5.'), l.")7, pi. <S, fig. 2 — in, in
K. D. Vid. Sel.sk. Skr., Ser. (J, vol. 3, pp. 287, 291, 393.
Ill his very much larger species, Alcirona krehsii, out of seven examples Hansen
describes and figuri's a male, i)robably aijidt, 92 nnn. long, and a great non-ovigorous
female 18 mm. long. These are contrasted iu shape by the circumstance that tlio
84—2
638 ON CRUSTACEA BROUGHT BY DR WILLEY FROM THE SOUTH SEAS.
peraeon segments of the female are much narrower and, except the first, much longer
than those of the male. Of Alcirona insulari^ he had at command three specimens,
but these comprised no female, were not fully grown males, and were not all exactly
alike. They ranged in size from 3'7 to o"l mm., the longer ones having more joints
iu the antenual flagella, the telson distally more narrowed, and a greater hirsuteness
on the hinder half of the body. The shajje is likened to that of Cirolana parva, so
that these joung males, as also the figure shows, are in agreement with the male of
Alcirona krebsii in general appearance.
Two specimens were obtained by Dr Willey which do not seem easily separable
from Hansen's insiilaris. The}' are a male and a female, both adult, the former about
5 mm., the latter about 4 mm., in length. But the singular thing is that here it is
the male which has the segments of the peraeon much narrower and longer than
those of the female, just the reverse of what is found in Alcirona krebsii, and this
singularity remains whether the identification of the specimens with Hansen's species
be valid or not. There are certain differences to be noted, but these may well be
attributed to the maturity of the specimen.
The male differs from the female not only in the proportions of the peraeon,
but also in the pleon, the penultimate segment of which has a small projecting tubercle
in the middle of the hind margin while the triangular terminal segment is distally
very narrow, its serrate sides and spinulose truncate point being girt with plumose
setae. Down the centre there is a low ridge. On the seventh joint of the first
guathopod the four teeth are less erect than in Hansen's figure, lying in the direction
of the nail and being successively larger. Those of the second guathopod are a little
smaller but have otherwise the same character. Also in both gnathopods there are
six or seven little nodular teeth fringing the inner margin of the sixth joint. The
spines of the fourth joint in the gnathopods and also in the first peraeopods are very
stout. The second pair of pleopods are just like those figured by Hansen for the male
of Alcirona krebsii.
The female specimen had the marsupium filled with twelve large oval hardened
eggs, two of them brown, the rest pale. The general habit was like that of the young
male, but with the telson distally more narrowed than in Hansen's figures. The first
antennae have the first joint thickened at the base, much shorter than the second, the
thu'd joint in one antenna equal to the first joint of the five-jointed flagellum, in the
other antenna shorter than that joint. The second antennae in both sexes have 20 — 21
joints in the flagellum. In the maxillipeds of the female the broad vibratory lamina of
the second joint reaches slightly beyond the rather narrow palp. In the gnathopods the
seventh joint has two minute teeth followed by a longer one adjoining the nail, much
like the formation in the first peraeopods of the male. In the female all the peraeopods
have the finger with its margin simple except for a prominence at the base of the nail.
Habitat. Blanche Bay, New Britain, 1.5 — 20 fathoms.
^ga niultidigita, Dana, 6'3 mm. long, from Balabac Passage, north of Borneo, is
certainly an Alcirona, and may possibly be identical with the present species. In the
terminal segment it agrees fairly with the account of the male pleon above given, but
the size and shape of the body differ ; the proportions of the first antennae, and the
ON CRUSTACEA BROUGHT BY DH WILLKV FROM THE SOUTH SEAS. 639
teeth on thu tiugcrs ut the giiatliopods caiiiKjt bu iLcuiiciKil with my duscriptions, and
there is no tubercle on the penultimate segment of the pleon.
Fam. Cv.mothoidae.
liSOO. Gymothoidae, Hansen, ' Cirolauidae,' Vidensk. 8elsk. Skr., Ser. (i, Natiuv. Afd.,
vol. 3, pp. 316, 406.
1895. Cyviothoimie, Hansen, Isop. Cumae. Stomat. Plankt(jn-Exp., pp. 12, l-t.
1897. Cymothoinae, Hansen, Bull. Mus. Conip. Zool. Harvard, vol. 31, no. .5, p. 101.
1S99. Cymothoidae, Harriet Richardson, Proc. U. S. Mus., vol. 21, p. 828.
1900. Cymothoidae, Harriet Richardson, The American Naturalist, vol. 34, p. 219.
1900. Cymothoidae, Stcbbing, Marine Investigations of South Africa, Crustacea,
p. 55.
Having so recently discussed the fuller synonymy of this group, I may here limit
myself to repeating that the family in Hansen's acceptation includes the Anilocridae,
Saophridae, and Cymothoidae of the Monographia Cymothoarum by Schiiidte and
Meinert.
Gen. Anilocra, Leach.
1818. Anilocra, Leach, Diet. Sci. Nat., vol. 12, pp. 348, 350.
1818. Canolira, Leach, Diet. Sci. Nat., vol. 12, pp. 348, 850.
1881. Anilocra, Schiodte and Meinert, Mon, Cymothoarum, Nat. Tidsskr., Ser. 3,
vol. 13, p. 100.
1899. Anilocra, H. Richardson, Proc. U. S. Mus., vol. 21, p. 830.
1900. Anilocra, H. Richardson, The American Naturalist, vol. 84, p. 220.
1900. Anilocra, Stebbing, Marine Investigations of South Africa, Crustacea, p. 56.
Other references are supplied in the last cited memoir. Canolira has what is
called page precedence over Anilocra, from which Leach distinguished it by the single
feature that the rami of the uropods are almost equal, of moderate length, with the
inner ramus slightly the longer, while in Anilocru the rami are unequal, elongate, the
outer the longer. This difference being untenable as a generic character, the two genera
coalesce, and Canolira as standing hrst would naturally have become the generic name,
but the only species which Leach assigns to it, Canolira rissoniana, of unknown habitat,
seemingly has not been identified, so that the name Anilocra, with wiiidi Leach con-
nected some well-tuscertained species, has properly been adopted.
Anilocra dimidiata, Bleeker.
1857. Anilocra dimidiata, Bleeker, Crust. Ind. arch., p. 31, jjI. 2, tig. 10 — 10(/.
1880. Anilocra dimidiata, Aliers, Ann. Nat. Hist., Ser. 5, vol. 5, p. 462.
1881. Anilocra dimidiata, Schi6dt(! and Meinert, Mon. Cymothoarum, N.ii. Tidsskr,,
Ser. 8, vol. 13, p. Ill, pi. 8 (15), fig.s. 5, 6.
640 ON CRUSTACEA BROUGHT BY DR WILLEY FROM THE SOUTH SEAS.
Schibdte and Meinert, from whom I have borrowed the reference to Bleaker,
place side by side leptosoma and dimicUata as two species of Anilocra- having in
common geniculate first antennae, side-plates not carinate and the fingers of the first
four pairs of legs inflated in the middle. The ovigerous female of A. leptosoma is
described as attaining a length of 33".5 mm., with a body long elliptic, three or four
times longer than broad (almost as 10 : 3). The o\dgerous female of A. dimidiata
has assigned to it a length of 265 mm., with a body elliptic, scarcely three times
as long as broad (20 : 7). But Miers gives the length of this species as an inch
and a third, therefore practically as long as A. leptosoma. A specimen in Dr Willey's
collection is 35 mm. long by 10 mm. broad. Consequently the shape is not a dis-
tinguishing character between the two species. The points on which I rely for
identifying Dr Willey's specimen with A. dimidiata, as described by Schiodte and
Meinert, are the following. It has the ' front margin of the first peraeon segment
manifestly trisinuate, with the lateral sinuses much deeper than the middle sinus,'
the sides of the fifth pleon segment 'deeply, angularly incised.' and the terminal,
obscurely carinate, segment not produced into an apical point. To these features
may perhaps be added the character that the uropods extend very slightly beyond
the telson. According to Schiodte and Meinert they do not quite reach the end
of the telson in A. dimidiata, while in A. leptosoma the inner ramus reaches far
beyond it. Koelbel in his Anilocra alloceraea speaks of the two rami as nearly
equal and both reaching somewhat beyond the telson. His species is confidently
identified with A. leptosoma by Schiodte and Meinert, and doubtfully by Miers.
The latter author {he. cit. p. 463) says, ' Bleeker, it may be observed, notes that
the uropoda in A. leptosoma do not reach beyond the extremity of the terminal
post-abdomiual segment ; in his figure, however, they are represented as distinctly
longer than this segment, in this particular agreeing both with Kolbel's de.scription of
A. alloceraea and with the specimen before me.' It may be remarked that the
extension of the uropods in relation to the telson is not always easy to determine,
as the appearance varies, according as the rami are directed inward or outward, and
according to the amount of flattening to which the specimen is exposed.
In the specimen 35 mm. long the young could be perceived through the partial!}'
pellucid plates of the marsupium. These showed the head between the dark eyes
thickly covered with dark stellate markings. A second specimen measures 22 mm. by
6'25 mm., and a third 15 mm. by about 4"5 mm.
Habitat. One label in the bottle with these specimens read, ' Cymothoa off fish
called Losilili. Karuana, Nov. 1895,' the other, ' D'Entrecasteaux group. British New
Guinea.'
Gen. Renocila, Miers.
1880. Renocila, Miers, Ann. Nat. Hist., Ser. 5, vol. 5, p. 464.
1884. Renocila, Schiodte and Meinert, Mon. Cymothoarum, Nat. Tidsskr., Ser. 3,
vol. 14, p. 414.
According to Miers, 'this genus, in all its characters, is most nearly allied to
Anilocra, from which it is distinguished by its broad non-inflexed front, the greatly
l^roduced postero-lateral angles of the three posterior thoracic segments, and the greatly
ox CRUSTACEA BROUGHT I'.Y DR WU.LEY KROM THE SOUTH SEAS. 641
dilated superior antennae.' According to Schiiidte and Meinert, it is not to Anilocru,
but to Nerocila that it stands nearest, though easily distinguislicd from that genus
by tlie character of the antennae, and many other points. While Miers includes in
the generic character ' the front not produced inferiorly, so as to conceal the bases
of the antennae,' the other authoi-s give ' frons prosiliens, declivi.s, fornicata, scapos
antennarum magnani partem obtegens.' This concealment, in fact, as often, only applies
to the dorsal view. Miers, in describing the type species, Renocila uvata, a.ssigns to
the head a 'straight anterior uiargin, which is inflexed, but not produced so as to
conceal the bases of the antennae,' and this i.s evidently more accurate than the
expression ' non-intlexed' in the generic account.
The mouth-organs are not discussed by the above-named authors. In respect to
the upper lip, the two maxillae, and to some degree the maxillipeds, these agree with
the figures of those parts in Savigny's Egyptian Crustacea, pi. 11, fig. 10, assigued
by Audouin to Anilocra cuvieri, Leach, but copied by Guerin, Iconogi-aphie, Crust.,
pi. 29, fig. 4, for his Canolira aecjijptiaca. Seeing that Gut^rin expressly says that
his species has the outer ramus of the uropods a little longer than the inner, and
Canolira is di.stinguished by Leacli from Ajiilocra by the single character that the
inner ramus is the longer, Guerin's species cannot be a Canolira, and that genus
appears to be unteuable for want of any real definition.
Renocila pehiophth.vlmi, u. sp.
Plate LXVIII B.
Head broader than long, front slightly inflexed, truncate with rounded angles ;
breadth abruptly increasing at the eyes. Body smooth ; peraeon broad, only slightly
convex, length of .segments in medio-dorsal line least in the seventh, greatest in the
first, which is nearly approached by that of the fourth ; obtuse-ended sidi^-plates of
second and third segments reaching beyond the lateral angles : those of the fourth
about, or not quite, level with those angles, those of the three following segments
successively much smaller and not nearly reaching the angles, which are successively
more produced backward, those of the last segment very obtuse and overlapping the
first three segments and nearly all the fourth of the abruptly narrowed, and in
(li.rsal view, parallel-sided pleon. The first segment of the pleon is rather longer
than any of the four following; all are obtusely angled medio-dor.sally : their side-
plates bend abruptly downward and then a little outward, being successively snuiUer,
and in the fifth segment not reaching thi> dor.so- lateral angles. The tennin.il segment
is flat, broader than long, strongly I'ounded exce])t at the base, so as to have an
almo.st circular appearance; its length ecpials that (jf the other pleon .segments
together.
Eyes rather obscure, small, distant, with about ten ocelli.
First antennae stout, not geniculate, the two btisal joints clearly distinct, the
third abruptly wider than the .second, the seventh much narrower than the sixth.
but still broader than long, the eighth minute.
Second anteimae quite concealed in dor.sil view, much shorter and narrower than
the first, consisting of seven joint.s, none very large.
642 ON CRUSTACEA BROUGHT BY DR WILLEY FROM THE SOUTH SEAS.
Upper lip with distal margin formiug four well separated blunt teeth or lobes.
Mandibles. Cutting edge broad, produced downward in a long blunt tooth or
process, not strongly chitinized, and showing above a tooth which seems to stand
free, but from the curvature of the plate has its bearings obscured ; palp is planted
rather far back, first joint very large, second smaller but still rather large, and
attached angularly below the apex of the first, third small, tipped with one or
more setules. On the left mandible this joint in our specimen carries four setules
and is narrower and less conical than on the right, but the differences may be
accidental.
First maxillae. The slender plate is tipped with four small curved spines.
Second maxillae. The apex is unequally divided between the little inner plate
and the broad outer, the former carrying one, and the latter two or three extremely
small hooked spines and others still smaller that are not hooked.
Maxillipeds. Second joint the largest, with very convex outer margin, the third
broadly tapering, the fourth small, bent inward and tipped with a little outward
pointing hooked spine.
The limbs of the trunk differ little in length. They have the second joint parallel-
sided, the fourth and fifth very short, the sixth with an almost circular apex over-
lapping the base of the finger, the finger geniculate in the first four pairs, in the
last three simply but strongl}' uncinate, not longer than the sixth joint.
The second pleopods have the stiliform process shorter than the outer ramus,
and the much larger inner ramus has a faint transverse suture above the middle.
The uropods have the peduncle slightly produced at the inner apex, the rami
slightly curved, blunt-ended, the outer the longer, reaching nearly the end of the
tenninal segment of the pleon.
Length, 12 mm., breadth, 6'2.5 mm.
Habitat, Lifu, parasitic on Periophthalmus, whence the specific name.
Renocila ovate, Miers, attains a length of 24 mm. Schiodte and Meinert say that
it has the terminal segment much longer than broad, once and a half as long as
the other pleon segments united. Yet they also say that it is transversely suboval,
and Miers says that it is almost semicircular in outline, giving a figure in which
it is decidedly broader than long. The postero-lateral angles of the seventh peraeon-
segment reach its base, which is not the case in the other two species of the genus.
Eenocila indica, Schiodte and Meinert, attains a length of 18 mm., and has the fingers
of the trunk-limbs elongate, those of the first pair being much longer than the sixth
joint.
Gex. Meinertia, Stebbing.
1893. Meinertia, Stebbing, History of Crustacea, Internat. Sci. Ser., vol. 74, p. 354.
1899. Meinertia, H. Richardson, Proc. U. S. Mus., vol. 21, p. 829.
1900. Meinertia, H. Richard.son, The American Naturalist, vol. 34, p. 220.
1900. Meinertia, Stebbing, Marine Investigations of South Africa, Crustacea, p. 57.
The name of this genns takes the place of Ceratothoa, Schiodte and Meinert,
which is distinct from the earlier Ceratothoa of Dana.
ON CRUSTACEA BROUGHT B^' DK WILLEY FROM TUK SOUTU SEAS. 643
Meinertia gaudichaudii (Milne-Edwards).
1840. Cymothoa Gaudichaudii, Milne-Edwards, Hist. Nat. Crust., vol. 3, p. 271.
1868. Ceratothoa rapax, Heller, Reise Novarae, Crust., p. 14G, pi. 12, fig. 17.
1883. Ceratothoa Gaudichaudii, Schiodte and Meiucrt, Naturhist. Tidsskr., Scr. 3,
vol. 13, p. 33.5, pi. 13(20), figs. 11—1.5, pi. 14 (21), figs. 1—5.
1893. Meinertia Gaudichaudii, Stebbing, History of Crustacea, Internat. Sci. Ser.,
vol. 74, p. 354.
1899. Meinertia gaudichaudii, H. Richardson, Proc. U. S. Mas., vol. 21, p. 829.
Two specimens labelled 'Parasites from floor of nioiilh of Tetrodon Pauaieti,' agree
very closely with the clescription and figures of the adult male of this species given
by Schiodte and Meinert. Yet there are some small differences. In the first antennae
there are eight joints instead of seven. The front margin of the first peraeon segment
is in the middle more broadly and deeply concave. In the last peraeopod the hind
margin of the second joint is much less convex than in the figure. The fifth segment
of the plcon has its hind margin bisinuate rather than deeply quadrisinuate. The
terminal segment is not slightly carinate, nor is it manifestly longer than the rest
of the pleon segments combined. The inner branch of the uropods is a little shorter
instead of longer than the outer, and both branches are apically blunt. The authors
quoted give the length of the adult male as 13 — 32 mm., and, as the specimens here
described have a length of only 13 nnii., it is probable that some of the ditierences
mentioned would have disap|)eared with increase of size. It must, however, be observed
that the terminal segment in ISchiiidte's and Meinert's own figure is shorter than the
other pleon segments combined. Several species in the genus have eight-jointed first
antennae, but none of these combine the obtusely fronted head with the subtrapezoidal
terminal segment of the present .species. Cinusa tetrodontis, Schiodte and Meinert, of
which it is said that the female with the male has prett}' often been taken in the
mouth or jaws of Tetrodon Honckenii Bl., has the fir.st anteiniae seven-jointed, the
second ten-jointed, and the head, sides-plates, and terminal segments differently shaped
from those of Meinertia gaudichaudii.
Fam. Ri'HAEROMIDAE.
1847. Sphaeromidue, White, List of Crustacea in the British Museum, p. 102.
1900. Sphaeromidae, Stebbing, Proc. Zool. Soc. London, p. .552.
I have so recently given numerous references to authorities on this family, that
it seems unnecessary here to repeat them.
Gen. Cilicaea, Leach.
1818. Cilicaea, Leach, Diet. Sci. Nat., vol. 12, pp. 341, 342.
1825. Cilicaea, Desmarest, Consid. gen. Crust., p. 295.
1838 ? Cilicaea, Gu^rin-M^neville, Iconographic Regne Auim., Crust., pi. 30 {Cilicea
in text, p. 30, with correction of erroneous numbering on plate).
w. V. 86
644 ON CRUSTACEA BROUGHT BY DR WILLEY FROM THE SOUTH SEAS.
1840. Nesea (part), Milne-Edwards, Hist. Nat. Crust., vol. 3, p. 218.
1881. Cilicaea, Haswell, Proc. Linn. Soc. N. S. Wales, vol. 5, p. 475, vol. 6, p. 2.
1882. Cilicoea, Haswell, Catal. Australian Crust., p. 29.5.
1886. Cymodocea, Beddard, Challenger Reports, vol. 17, Isopoda, p. 145.
189.3. Cilicaea, Stebbing, History of Crustacea, p. 364.
1900. Cilicaea, Harriet Richardson, The American Naturalist, vol. 34, pp. 222,
224.
In a s}Tioptic table of the Sphaeromidae Miss Richardson distinguishes Cilicaea
as having 'outer branch of the uropoda not rudimentary,' 'only the external branch
of the uropoda projecting and exposed ; outer branch incapable of folding under
inner' ; ' all the thoracic segments of equal length. Penultimate abdominal segment
in male generally produced in spine. Terminal segment excavated with or without
median lobe.'
If, however, the figures given by Desmarest and Guerin of the t}-pe species
can be trusted, that has the first peraeon segment decidedly longer than the others,
nor will equality of the peraeon segments apply to any one of the six species
described by Professor Haswell. Mr Beddard considers that both Cilicaea and Nesea
are synonyms of Cymodoce, but while so little is known as at present about the
mouth-organs of the numerous species, speculation as to their generic position seems
useless.
Cilicaea tenuicaudata, Haswell.
1881. Cilicaea tenuicaudata, Haswell, Proc. Linn. Soc. N. S. Wales, vol. .5, p. 475,
pi. 17, fig. 2.
1882. Cilicaea tenuicaudata, Haswell, Catal. Australian Crust., p. 295.
Having had no opportunity of examining any other specimen of this genus it
is only provisionally that I assign the solitary one in Dr Willey's collection to
Professor Haswell's species. It agrees in almost all respects remarkably well with his
description and figures, but under the microscope the tuberculation, especially of the
pleon, is scarcely to be called obscure; the first antennae have 10 — 11, and the
second 14 — 15, joints in the flagella (whereas Haswell says, flagella of antennae each
with about ten articuli) ; and the long ramus of the uropods has the apex oblique,
more as in figure of C. crassicaudata, Haswell, not bifurcate as in both figure and
description of C. tenuicaudata. The eyes are large and deeply let into the front of
the large first segment of the peraeon. The long, apically bifurcate process of the
pleon is ventrally clothed with hair or stiff but fine setae. The length, 7'5 mm.,
agrees nearly with the five-sixteenths of an inch given by Haswell.
At the tip of one uropod is a little foraminifer. The apices of both uropods
and of the pleon process were grouped about this object, as if engaged either in
securing it or attempting to dislodge it. In this position the uropods seemed as if
they were bifurcate, but not so when separated.
Habitat. Blanche Bay, New Britain, 40 — 50 fathoms. Taken by trawl.
ON CRUSTACEA BROUGHT BY DR WILLEY FUO.M TUK SOUTH SEAS. 645
ONISCOIDEA.
ISOPODA TERKESTUIA.
1825. Oniscides, Latreille, Fam. Nat. du llegiic Animal, [>. 297.
18.5.S. Oniscoidea (part), Dana, U. S. Expl. Exp., Crust., p. 713.
1876. Oniscoidea, Miors, Crust. X. Zealand, p. 'M.
1882. Oniscoidea, Sars, Christiaiiia Vidensk. Forh., No. 18, p. .58.
1885. Isopoda terrestma, Budde-Luiid, Monograph.
1893. Oniscoidea, Stebbing, History of Crustacea, p. 420.
1898. Oniscoida, Sars, Crustacea of Norway, vol. 2, pt. 9, p. 153.
1900. Oniscoidea, H. Richardson, The American Naturalist, vol. 34, p. 301.
1900. Oniscoidea, Stebbing, Proc. Zool. Soc. London, p. 563.
This group answers to the Cloportides of Latreille and Milne-Edwards, and under
various headings has been in recent years largely illustrated by the well-known writings
of M. Adrien Dollfus. Buddc-Lund in 1885 uses the term Oniscoidea in a much
restricted sense, as the second section of his family Onisci.
Fam. Li(iiii)AE.
1885. Licjiae (part), BuJdo-Lund, Isopoda terrestria, p. 242.
1893. Ligiidae (part), Stebbing, History of Crustacea, p. 420.
1898. Ligiidae, Sars, Crustacea of Norway, vol. 2, pt. 9, p. 155.
1900. Ligiidae, H. Richardson, The American Naturalist, vol. 34, p. 306.
In 1898 Sars restricted this family by separating from it several genera wiiich
he allotted to a new family Trichoniscidae. With rapidly increasing knowledge of
the terrestrial isopoda some changes in classification cannot f;iil to ensue, although
the discoveries which necessitate the separation of group.s are always liable to be
counterbalanced by others which tend to reunite them.
Gen. Ligia, J. C. Fabncius.
1798. Ligia, Fabricius, Supplenientum Ent. Syst., p. 301.
1833. Ligia, Brandt, Bull. Soc. Nat. Moscou, vol. 6, p. 171.
1853. Lygia, Dana, U. S. Expl. Exp., Crust., p. 716.
1879. Ligia, G. M. Thomson, Trans. N. Zealand Inst., vol. 11, p. 232.
1885. Ligia, Buddc-Lund, Isopoda terrestria, p. 258.
1893. Ligia, Dollfus, Feuille dcs Jeunes Naturalistes, Sor. 3, Annde 24, No. 273.
1S99. Ligia, Chilton, Ann. Nat. Hist., Ser. 7, vol. :), p. IHT.
Buddc-Lund cites the spelling Lygia also from Roux, 1828. Dollfus distinguishes
this genus as neither terrestrial nor marine, but maritime, as living not in the sea
but on its lips. Chilton's paper is on the se.xual characters of Ligia oceanica. Budde-
Nund referring, confessedly at seconil liaml. to Thomson's Ligia quadrata, gives the
habitat as Australia. It should ]»• Duncdiu, New Zealand.
85—2
646 ON CRUSTACEA BROUGHT BY DR WILLEY FROM T^E SOUTH SEAS.
LiGiA viTiENSis, Dana.
1853. Lygia vitiensis, Dana, U. S. Expl. Exp., Crust., p. 741, pi. 49, fig. 5 a, h.
1885. Ligia vitiensis, Budde-Lund, Isopoda terrestria, p. 271.
Dana's single specimen from 'the Feejees' was 'mutilated in its last abdominal
segment, besides wanting the stylets and antennae.' In all Dr Willey's specimens the
stylets are unfortunately missing. The eyes are large, widening outward, the space
between them more than half the horizontal length of the eye, not less than half as
in Dana's Ligia hawaiensis. The second antennae in natural position reach back to the
beginning of the pleon ; the last joint of the peduncle is considerably longer than the
penultimate ; the flagellum is rather longer than the peduncle, with 28 to 30 joints.
In the last two pairs of trunk legs, that is, the fourth and fifth peraeopods, there is
a tuft of hairs on the hind margin at the base of the double unguis. The terminal
segment of the pleon has the postero-lateral angles rather long, acute, but otherwise
its apical border is very unlike that described and figured by Dollfus for Ligia
exotica, Roux, the extremity being as Dana says ' very low, triangular' and the inter-
mediate angles being, as his figure shows, quite blunted down. This margin and the
sides of the segment carry minute spinules, of which two flank the little apical
emargination or notch. Dana speaks of the surface of the body as quite smooth,
but there are scattered hairs in our specimens. Colour, the usual diversified iron
grey.
Length, from front of head to end of telson, 13 — 17'5 mm., only a single
specimen attaining the latter dimensions. For Ligia exotica Budde-Lund gives 20 — 30
( — 35) mm. for the length.
Habitat. Matadona, China Straits, British New Guinea. ' From face of cliff, with
fresh water species, far above tide-mark.'
Fam. Oniscidae.
1885. Onisci (Section II.), Budde-Lund, Isopoda terrestria, p. 75.
1893. Oniscidae, Stebbing, History of Crustacea, p. 426.
1898. Oniscidae, Sars, Crustacea of Norway, vol. 2, p. 169.
1900. Oniscidae, H. Richardson, The American Naturalist, vol. 34, p. 302.
Budde-Liind's family Onisci comprises two sections, the Armadilloidea and Onis-
coidea, corresponding to the two families Armadillidiidae and Oniscidae. In the
latter group he includes a gen-us Oniscus, which he divides into five subgenera,
Oniscus, Philoscia, Alloniscus, Lyprohius, Scyphax. It is by most writers, I believe,
thought more convenient to regard all these as independent genera. Budde-Lund
himself assigns twenty-three species to Philoscia, and several have been added since
his book was published.
Gen. Philoscia, Latreille.
1804. Philoscie (probably misprint for Philoscia), Latreille, Hist. Nat. Crust, et
Insectes, vol. 7, p. 43.
ON CRUSTACEA BROUGHT BY DR WILLEY FROM THE SOUTH SEAS. 647
1810. PInloscia, Latreille, Consid. gen. Crust., Arachnides, Insectes, p. 110.
1813. Philoscia, Leach, Edinb. Encycl. (Art. Crustaceology), vol. 7, p. 406.
1825. Philoscia, Desmarest, Consid. gen. Crustaces, p. 318.
1833. Philoscia, J. F. Brandt, Bull. Soc. Nat. Moscou, vol. 6, p. 182.
1840. Philoscia, Milne-Edwards, Hist. Nat. Crust., vol. 3, p. 1G3.
1868. Philoscia, Bate and Westwood, Brit. Sess. Crust., vol. 2, p. 448.
1880. Philoscia, Harger, Rep. U. S. Comm. Fisheries, pt. 6 for 1878, p. 30.5.
1885. Philoscia, Budde-Lund, I.sopoda terrestria, pp. 201, 207.
1893. Philoscia, Stebbing, History of Crustacea, Internat. Sci. Ser., vol. 74, p. 430.
1897. Philoscia, Dollfus, Feuille des Jeunes Naturalistes, Ser. 3, 27'= Annee, No. 317
(Tableau icon, des Philoscia d'Europe).
1898. Philoscia, Sars, Crustacea of Norway, vol. 2, p. 172.
1900. Philoscia, Harriet Richardson, The American Naturalist, vol. 34, p. 303.
In this genus the antennae are comparatively smooth, the first maxillae have
several spines on the apex of the outer plate, and the inner ramus of the uropods
is slender, these being characters in which it differs from the genus next to be dis-
cussed.
Philoscia gracilis, Budde-Lund.
1879. Philoscia gracilis, Budde-Lund, Prospectus Crust. Isop. terrestrium, p. 2.
1885. Philoscia gracilis, Budde-Lund, Isopoda terrestria, p. 220.
A single specimen, without antennae or uropods, appears to belong to this species,
which I only know from the second work above cited. 'The fifth segment of the
peraeon almost entirely black-brown' among the others that are much marbled with
white is rather a striking characteristic. The length of the specimen, however, is only
about 6 mm., whereas Budde-Lund's from the Island of Upolu was 10 mm.
Habitat. Lifu, Loyalty Islands.
Philoscia truncata, Dollfus.
1898. Philoscia truncata, Dollfus, Zool. Ergebn. Niederlandisch Ost-Indien, vol. 4,
p. 376, pi. 15, fig. 23, and in text fig. 23 o, 23 6.
The .shape and colouring of the specimens agree very exactly with the figure on
Dollfus' plate, the seventh segment of the peraeon having the postero-lateral angles
notably truncate, with the whole of each corner pale in contrast with the dark hue
otherwise prevailing. A specimen 8 mm. in length, which is the length given by Dollfus
for specimens from Celebes and Flores, has the caudal segment apically .sub-obtuse ;
in a smaller specimen, only 5 mm. long, this segment would rather be described as
obtuse at the apex.
Habitat. New Britain.
648 ON CRUSTACEA BROUGHT BY DR WILLEY FROM THE SOUTH SEAS.
Philoscia lifuensis, n. sp.
Plate LXIX B.
The surface scabrous with minute, sparsely planted hairs, some of them occurring
between the lenses of the eyes. Bent part of the occiput forming a dark border to
the head in a ventral view. First three segments of the peraeon with the hind margin
very slightly sinuous, last segment the longest, none with acute angles, all pale in
front, and darkly marbled behind. Pleon abruptly narrowed, short, the caudal segment
much broader than long, very obtuse at the apex.
Eyes with several ocelli, probably about sixteen. Second antennae missing.
First maxillae with the outer plate rather narrow, apically bent, the apex carr}'ing
eight spines, the outermost the largest, several bifid, the setulose fringe of the concave
distal part of the outer margin having its little setules much longer at the extre-
mities than in the middle portion. The maxillipeds have a minutely spinulose surface.
Uropods. Peduncle reaching be3"ond the telson, channelled on the outer edge, in-
termediate in length between the two lanceolate rami, which are attached on a level,
the slender inner one three-fifths of the length of the more robust outer one, both
spinose. Rami and peduncle alike are microscopically fi'inged at the inner margin.
Colour of dark portions blackish or purplish grey.
Length, 42 mm., breadth, l"^ mm. Philoscia lueberi, Dollfus, 1898, from Sumatra, is
5'5 by 24 mm., has no dark band on the middle segment of the peraeon, the dark bands
on the first three segments in the front instead of at the back, and, judging by the
figure, has a long pleon, not a short one. In other respects it shows agreement with
the form here described.
Habitat. Lifu, Loyalty Islands. Specific name from place of capture.
Paraphiloscia, n. gen.
In general agreement with Plnloscia, but distinguished by the spinose second
antennae, first maxillae with only three apical spines on the outer plate, and the
uropods with laminar inner ramus, and each ramus terminating in a pencil of setae.
The spinose somewhat geniculate antennae and the long setae at the tips of the
uropods recall the features of Trichoniscus, but connexion with that genus is excluded
by the mouth-organs, the eyes with numerous ocelli and the telson not truncate.
Paraphiloscia stenosoma, n. sp.
Plate LXIX c.
Head with deflexed front. Peraeon narrow, nearly parallel-sided, the earlier seg-
ments a little shorter and broader than the hinder ones. Pleon abruptly narrower,
slightly tapering ; anterior margin of the first segment seen through the overlapping
hind margin of the seventh peraeon segment gives the appearance of a segment too
many. Caudal segment twice as broad as long, subacutely triangular.
Eyes with about twenty-one ocelli. First antennae minute, first joint subequal to the
ON CRUSTACEA BROUGHT BY DR WILLEY FROM THE SOUTH SEAS. 649
second and third united. Second antennae with the three joints of the flagellum sub-
equal, and together equal to the last joint of the peduncle, which is slightly longer
than the penultimate, and considerably longer than the antepenultimate. Upper lip
with convex distal margin. First maxillae with two short thick plumose setae on the
inner plate, three spines on the apex of the outer, one of them considerably stouter
than the other two ; the outer margin of the apical part is very feebly fringed, the
inner margin carries some setules. The second maxillae have the apex divided between
a very small process and a rather broad plate. In the maxillipeds the epipod is more
than two-thirds as long as the large second joint, which is capped by a small apically
rounded almost unarmed plate ; the palp is a little shorter than the plate, though
reaching beyond it with its curved apical seta.
The legs are rather spinose, with the second joint not very large, but the third
to the fifth stout, the sixth being abruptly narrower; the finger is slender, with two
setules on the inner margin.
Uropods. The peduncle reaches little beyond the caudal segment, and is equal
in length to the finely fringed inner ramus, which is not quite twice as long as
broad, though seen edgewise it gives a very different impression. It is inserted a little
in front of, and is about three-fifths as long as, the lanceolate outer ramus.
Colour, uniform light brown.
Length, 3"60 mm., breadth, about 1'1.5 mm.
Habitat. New Britain.
" Oniscus ? angustus," Dana, from Tierra del Fuego, shows some resemblance to
this species, for which I had at first chosen the same specific name. Dana's specimen,
which was more than twice as long, had lost the antennae and uropods,
Fam. Armadillidiidae.
1885. Onisci (Section I.), Budde-Lund, Isopoda terrestria, p. 14.
1893. Armadillididae, Stebbiug, History of Crustacea, p. 432.
1898. Armadillidiidae, Sars, Crustacea of Norway, vol. 2, pt. 11, p. 187.
1900. Armadillididae, H. Richardson, The American Naturalist, vol. 34, p. 305.
Brandt in 1833 divides his group Armadillina into two sections, the first Arma-
dillidia, containing only his genus Armadillidium ; the second Cubaridea, containing
his two genera, Cubaris and DiploexocJms. Both sections are united in the present
family.
Gen. Cubaris, Brandt.
1833. Cubaris, J. F. Brandt, Bull. Soc. Nat. Moscou, vol. 6, p. 189 (Conspectus
Monograpliiae Crustaceorum Oniscodorum Latreillii).
1833. Armadillo, Brandt, ibid., p. 191.
1833. Diploexoclms, Brandt, ibid., p. 192.
1840. Armadillo, Milne-Edwards, Hist. Nat. Crust., vol. 3, 177.
1840. Diploea^ochus, Milne-Edwards, ibid., p. 180.
1840? Pentheus, C. L. Koch, Deutschlands Crustaceen, Myriapoden und Arachniden,
Heft 34, No. 1 (Herrich-Schaffer, Heft 180, No. 1).
650 ox CRUSTACEA BROUGHT BY DR WILLEY FROItf THE SOUTH SEAS.
1843. Armadillo, Kraiiss, Die Siiclafrikanischen Crustaceen, p. 63.
1847. Armadillo, White, Crust. Brit. Mus., p. 100.
18.53. Armadillo, Dana, U. S. Expl. Exp., vol. 13, p. 71.5.
1853. Spherillo, Dana, ibid., pp. 71.5, 719.
1853. Diploexochus, Dana, ibid., p. 715.
1859. Pyrrioniscus, Kinahan, Proc. Dublin Univ., vol. 1, p. 199.
18(58. Spherillo, Heller, Reise der Novara, Crust., p. 134.
1876. Armadillo, Miers, Catal. Crust. New Zealand, p. 94.
1876. Cubaris, Miers, ibid., p. 95.
1876. Spherillo, Miers, ibid., p. 96.
1877. Cubaris, Miers, Proc. Zool. Soc. London, p. 664.
1877. Orthonus, Miers, ibid.
1879. Armadillo, Budde-Lund, Prospectus Isopodam terrestrium.
1885. Armadillo, Biidde-Luud, Isopoda terrestria, pp. 15, 50, 282.
1887. Armadillo. Dollfus, Bull. Soc. detudes sci. de Paris (Crust, isop. terrestres).
1893. Cubaris, Stebbiug, History of Crustacea, Internat. Sci. Ser., vol. 74, p. 432.
1898. Armadillo, Dollfus, Zool. Ergebnisse Niederlandisch Ost-Indien, vol. 4, p. 358.
1898. Cubaris, Sars, Crustacea of Norway, vol. 2, p. 188.
1900. Cubaris, Harriet Richardson, The American Naturalist, vol. 34, p. 305.
This genus now contains so large a number of species that its subdivision would
be a matter of convenience. In the sj-nonymy there are several names inviting
employment for such a purpose. Unfortunately they were introduced before the
necessities of the case were well understood, so that in general they are rather a
stumbliug-bloek than an assistance. The use of the name Cubaris itself requires
vindication, especiallj' as it has been discarded both bj- Gustav Budde-Lund and by
Adrien Dollfus, two writers to whom science is so deeply indebted for extended and
more intimate acquaintance with the land Isopoda of the world. The name Armadillo
to which those authors give the preference was introduced by Latreille in 1804,
Hist. Nat. des Crustaces, vol. 7, p. 47, with the three species, vulgaris, variegatus,
maculatus. But by Budde-Lund the first two of these are included in the genus
Armadillidium, and the third is considered not to be an isopod at all. All three
are referred to Armadillidium by Dollfus. For these authorities, therefore. Armadillo
ought to disappear. They, like Brandt, base it upon a .species called Armadille des
boutiques by Dumeril in 1806 {officinalis, Desmarest, Consid. gen. Crust., p. 323,
officinarum, Brandt, loc. cit., p. 191), which, as Miers remarks, had not been described
when Latreille founded the geniis, so that, as Dumeril's Armadillo is generically
distinct from Latreille's, it falls to the ground as a preoccupied name. Latreille's
Armadillo, it is true, has precedence of Brandt's Armadillidium. But there is fair
reason for maintaining that Latreille's Armadillo is also preoccupied. The name was
used for a quadruped by Brissou in 1756, and the second enlarged edition of Brisson's
Regnum Animale in 1762 retains the name (see p. 23) in preference to the Dasypus
of Linnaeus. No doubt Brisson was not uniformly consistent in the use of the
recently-introduced binomial nomenclature, but he does not transgress it in any
of his seven species of Armadillo, except that he leaves the first without any
ON CRUSTACEA BROUGHT BY DR WILLEY FROM THE SOUTH SEAS. 651
specific name, as the Armadillo, which does not seem a bar! notion for indicating
the type of a genus. In any case it should be remembered that in the Stricklandian
Rules of 1842 Brisson is expressly mentioned as one whose defined genera had a title to
recognition, though the same indulgence was not to be extended to his species, even
when their designations ' are accidentally binomial in form.' Latreiile himself admits
preoccupation of the name in its French form, for in 1804, loc. cit., p. 63, when
establishing the myriapod genus Glomeris, he says that its formation is due to Cuvier,
'qui I'avoit nomme armadille' (Journ. d'hist. natur. tome ii. p. 27).
In the Regne Animal, vol. 4, p. 144, 1829, Latreiile assigns to his Armadillo
species belonging to both Brandt's genera Armadillidium and Armadillo, and it is
perhaps for this reason that Budde-Lund suggests that Latreille's original Ar-madillo
vulgaris may have been a collective species, from which Brandt had a right to extract
the later Armadillo officinalis as a type of the genus. But Latreiile defiuing
Armadillu in 1829 expressly says that the last segment is triangular, a shape of the
telson which excludes Dumeril's species, and entails the alternative of either substituting
Armadillo for Armadillidium or relinquishing Armadillo altogether.
It may be worth explaining, in regard to terminology, that Dollfus distinguishes in
the epistome three portions, first, the prosepistome, which is the upper boundary, in
this genus not unfrequently advanced in front of the cephalic 'front,' secondly, the
mesepistome, a middle region, in or on which lie the basal joints of the second
antennae, and thirdly, the metepistome, a lower transverse piece, flanked by lobes at
its upper corner, and bearing below it the labrum or upper lip.
CuBARis ciNCTUTUS (Kinahan).
Plate LXIX A.
1859. Pyrgoniscus cinctutus, Kinahan, Proc. Dublin Univ., vol. 1, p. 200, pi. 19,
fig. A—/:
1885. Armadillo translucidus, Budde-Lund, Isopoda terrestria, p. 290.
It is not without some hesitation that I identify the form here described with
Budde-Lund's species from Noumea, New Caledonia. The Danish author states that the
last joint of the peduncle in the second antennae is almost three times as long as
the flagellum, that the eyes are large, with rather large, subglobose ocelli, twenty or
more in number ; and that the colour is yellow, sprinkled with numerous black or
blackish spots and dots, being in particular black with confluent spots in the middle
of the fifth, sixth and seventh segments of the peraeon and at the sides of the third,
fourth and fifth segments of the pleon. In these respects his specimen is not in
agreement with Dr Willey's. But the agreement otherwise is so very close that
separation seems unadvisable. Budde-Lund in his Monograph, p. 46, .speaks of knowing
Kinahan's Pyrgoniscus by the description, but curiously leaves his readers without any
clue to the terms of it, which he had probably himself forgotten when describing
Armadillo translucidus in the appendix to his own work.
Kinahan's specimen from ' the Eastern seas ' was without the second antennae,
w. V. 86
652 ox CRUSTACEA BROUGHT BY DR WILLEY FROM THE SOUTH SEAS.
and we are not told that its mouth-organs were examined. Yet the placing of the
species in a new genus, if not quite tenable, was not without plausibility, for it
does not conform to one of the prominent characters assigned to the Armadillidiidae,
that of being contractile into a globular shape. The flattening of the head and tail
and sides prevents this, and the creature doubles up instead of rolling up, the side-
plates of the peraeon, especially those of the third and fourth segments, becoming con-
spicuously imbricated. In any future subdivision of the genus Cubaris, Kinahan's
Pyrgoniscxis should be taken into account.
In the following description the sentences in inverted commas are translated from
Budde-Lund.
Body " oblong oval, rather convex, subopaque, very minutely squamose and
punctate."
Eyes not very large, oval, with rather small ocelli, 15 — 17 iu number.
First antennae, as usual minute, thii'd joint a little longer than the second, with
some apical and subapical setules.
Second antennae. Last joint of peduncle about twice as long as each of the
three preceding joints and as the flagellum, " first joint of flagellum not half as long
as the second."
" Clypeus with large, oblong, roundly subrectangular lobes.'' This is the part called
by Dollfus the metepistome, which supports the labrum. Between the transverse
plate and the upper antennae, as shown iu the ventral figure of the head, there are
two outward-directed horns or lobes.
"Epistome [prosepistome of Dollfus] with its upper margin reaching much in
advance of the front, the middle faintly cleft and sub-bipartite by a longitudinal
suture. Front behind the plate of the epistome excavate in the middle." This laminar
expansion in front of the ' front ' by its great size and central cleft is the most
striking feature of the species.
The labrum or upper lip is transversely and narrowly oval. The lower lip has
two broad contiguous lobes, with a central conical inner plate.
The cutting edge of the mandibles is tridentate, powerful, darkly coloured like
the secondary plate, which is also strong ; near to these is what Sars describes as
a membranous hairy lappet and a group of setae, and a little more remote a re-
curved much subdivided seta.
The first maxillae have two short thick plumose setae on the inner plate, and on
the outer ten spines very unequal in length and thickness.
The second maxillae have .the apex divided into two lobes of very unequal
breadth.
Maxillipeds. The epipod is produced along the side of the large second joint, and
is slightly folded, and perhaps expanded at the apex. The small plate on the inner
apical margin of the second joint has a straight inner and convex outer margin, with
three spinules on its crenulate distal margin. It is fully as long as the small two-
jointed palp, which has two spinules on the transverse first joint and four on the
rounded triangular second.
"Segments of the peraeon with thin translucent side-plates. Side-plates of the
^
T RRS Del
•imbndge
Cubans cincLuLias.
[Kinahan]
Philoscia hfuf
Paraphiloscia steno^;ui:ia,i: y
ON CRUSTACEA BROUGHT BY DR WILLEY FROM THE SOUTH SEAS. 653
first three segments with a smallj inferior, tooth. Hind margin of the anterior
segments, especially the first, sinuated on each side, the hinder angles of the first
segment acute. The first segment longer and larger than the rest, in the middle
sub-bulbose, produced. Legs slender." By the ' small, inferior, tooth ' I understand
that to be intended which I have shown in the ventral figure of ^jer. s. 1 ; it is
produced backward at the sinuate point of the hind margin on either side. The
spinulation of the slender legs is sufficiently shown in the figures gjip. 1 and prp. 5.
" Side-plates of the third, fourth, and fifth segments of the pleon broad, rect-
angular, widening outward. Anal segment a little wider than long, narrowed in the
middle, having at the base a longitudinal fluting, the apex convexly truncate; the
peduncle of the uropods oblong, much longer than broad, at the apex a little narrowed-^
outer ramus small, inserted far from the apex in the incisure of the inner margin
of the peduncle ; inner rami much shorter than the anal segment." The longitudinal
stria or fluting may easily be overlooked in consequence of the colour variegation.
Colour in our specimens forming throughout a dark ground of brown and blackish
brown confluent spots, marbled with light markings, which in many parts, especially
on the sides, take the form of bright round or rounded dots ; under side and legs
pale, as is also the dorsal connecting skin between the segments shown in flexure.
Length, 10 — 12 mm., breadth, 5 — 6"25 mm. Budde-Lund gives length, 11-5 mm.,
breadth, 5o mm., height, 2'7 mm.
Habitat. Lifu, Loyalty Islands.
Miers (Proc. Zool. Soc. London, for 1877, p. 667), in his account of Gubaris gigas,
from Nicaragua, remarks that ' in the prominent anterior margin of the head, beneath
which the antennae are partly concealed, it resembles the genus Pyrgoniscus, Kinahan,'
and this genus, he adds, 'is nearly allied to Gubaris! In G. gigas the prominent
margin of the head is represented without any median fissure.
CUBARIS LIFUENSIS, n. sp.
Plate LXX a.
Body minutely squamose, with microscopic scattered hairs, otherwise smooth. Head
short and broad. Anterior margin of prosepistome rather convex, just forming a rim
to the front of the head. Finst peraeon segment with front and hind margins
strongly sinuous, anterior angles more acute than the posterior, lateral margins thin, in
front slightly curving ujjward, a slight fold passing, at a little distance from the outer
edge, from the front angle to a rounded ending some way short of the hinder angle.
This fold is scarcely to be seen without dissection. The hinder angles of all the
segments after the second are subquadrate, the second and third segments have a
duplicature ending roundly half-way down, not adjacent to the lateral margin.
The eyes have ocelli in varying number, 16 — 22.
In the second antennae the penultimate joint is longer than either of the two
preceding joints and slightly longer than the flagellum, which is two-thirds as long
as the ultimate, and has a first joint only a little shorter than the second.
The terminal segment of the pleon has its base shorter but much broader than
86— 2
G54 ON CRUSTACEA BROUGHT BY DR WILLEY FROM THE SOUTH SEAS.
its quadrate apical part, in which the sides are parallel, the distal margin a little
convex. The uropods (seen from below) are as broad as long, the much narrowed
distal part not closely filling the space between the terminal segment and its pre-
decessor; the inner rami reach the end of the terminal segment, their apical setules
being visible beyond it ; the conical outer rami reach the end of their own peduncle.
Colour mottled with brown and yellowish markings of an ordinary type. Length,
11—12 mm.
Habitat. Lifu, Lo}-alty Islands. Budde-Lund's Armadillo nignnus, of unknown
habitat, probably from the Cape of Good Hope, seems to approach this species nearly,
but it is described as very minutely grauular, with the ordinary tuberculosities
manifest on both sides of the bod\- : with 20 or more ocelli to the rather large eyes,
the terminal segment of the pleon scarcely broader than long, its rectangular apex
longer than broad, slightly carinate longitudinally ; the peduncle of the uropods much
longer than broad, the colour grey, or grey-black, with paler sides. The side-plates
of the (first) segment of the peraeon are said to be entire, thin.
CUBARIS DOLLFCSr, n. sp.
Plate LXX b.
This species makes the nearest approach to C. simplex (Dollfus), 1895, from North
Madagascar, but is distinguished from it by the longer rami of the uropods, and as
well by the anterior margin of the prosepistome as the apical of the caudal segment,
both of which are straight in the species compared.
The present species has the body only moderately convex, minutelj' squamose.
The prosepistome is well in advance of the front, broadly convex in the middle,
with subacute lateral lobes. Eyes prominent, with about 18 ocelli. Second antennae
have the penultimate joint of the peduncle much longer than either of the two
preceding joints; rest missing.
First segment of peraeon, with front angles more rounded than the hinder, a
little raised ; the duplicature represented by a long ridge remote from margin ending
in a small tooth ; the two following segments each have a short ridge, ending in a
tooth ; the sides of the sixth and seventh segments are very much broader than
those of the four preceding segments ; in the pleon the third, fourth and fifth segments
laterally widen outward.
The caudal segment is nearly as long as breadth at base, the basal part about
one-third of the length, the subquadrate apical part with sides very slightly con-
verging to a somewhat arched or obtuse-angled apex, thus supplying a form very
unusual in this genus. The segment is not quite flat, but slopes a little from the
middle to the sides.
The uropods with the narrow apical part do not completely fill the space between
the terminal and preceding segments. The inner rami reach the apical margin of the
caudal segment ; the outer are slightly curved, more than half as long as the peduncle,
being attached high up well within its margin, and reaching slightly beyond its
apex, though falling considerably short of the inner rami.
.Vi:--jDY. LOOLOOlCAi
Cubaris hfuensis,n sp.
C Tnlifi:^. r- <=^ r 1,,,^. „ sp C. zebricolor,n sp
Panaietis incamerata, n g. et sp
Bassettia c ^- - ^ ^^ sp
ON CRUSTACEA BROUGHT BY DR WILLEY FROM THE SOUTH SEAS. 655
The colour is brightly and on the whole symmetrically marbled with brown and
yellowish tints. The head is rather dark, with a transverse pale line in two portions
near the front and pale markings at the back. The legs are longitudinally striped
above with brown. Length, 11 mm., breadth, a little over 5 mm.
Habitat. Lifu, Loyalty Islands.
This interesting species is named in compliment to M. Aflrien Dollfus.
CuB.iRis OFFICINALIS (Desmarest).
1825. Armadillo officinalis, Desmarest, Consid. gen. Crust., p. 323.
1833. Armadillo officinarum, Brandt, Bull. Sec. Nat. Moscou, vol. 6, p. 191, pi. 4,
fig. 16, 17, 19.
188.5. Armadillo officinalis, Budde-Lund, Isopoda terrestria, p. 16.
1890. Aj-madillo officinalis, Dollfus, Soc. d'etudes sci, de Paris, Juillet, 1890, p. 5.
Although this species belongs to the countries surrounding the Mediterranean,
I feel obliged to assign to it specimens from the South Pacific, which are indeed
smaller in size and different in colour, but otherwise show no appreciable differences
that I can detect.
The body is very convex. The prosepistome is convex, scarcely passing beyond
the front. The eyes are small, with about 12 ocelli. The second antennae have the
earlier joints proportioned as in C. dollfusi, but the last joint of the peduncle and
the flagellum rather longer than in that species, the first joint of the flagellum half
as long as the second. The first segment of the peraeon has the lateral margins
sulcate and cleft at the distal end ; the duplicature is strongly expi-essed in the
second segment. The broad short telson has the wide basal part slightly longer than
the distal, of which the apical margin is very feebly convex. The uropods have a
broad apical margin closely fitting between that of the caudal and fifth segments ;
the inner rami not reaching the apex of the telson, the outer minute, only visible
above in a lateral pocket of the peduncle.
Colour mottled with dark brown and yellow, sometimes the brown prevailing,
diversified with pale spots.
Length of largest specimen rolled up, 8 nmi., unrolled it might be about 13 mm.
Habitat. Isle of Pines, South of New Caledonia.
Dollfus in 1892 describes 'a variety ex colore' from the Mount of Olives and
other Syrian localities as Armadillo officinalis, var. Syriaca.
CUBARIS LUNDl, n. sp.
Plate LXX c.
Body very convex, squamose ; each segment of the peraeon having on either side
of the back a little pit with a small elevation in the middle of it.
Both front and hind margins of the head concave in the centre. The pros-
epistome follows the frontal curve, from which it is separated by a narrow groove.
The eyes are not very prominent, composed of numerous (ab<iut 27) small ocelli.
656 ON CRUSTACEA BROUGHT BY DR WILLEY FROM THE SOUTH SEAS.
The second antennae are not elongate ; the short flagellum is about three-
quarters the length of the last joint of the peduncle ; of the three preceding joints
the middle one is the shortest, each of the others being about as long as the flagellum;
first joint of the flagellum half as long as the second.
The flrst segment of the peraeon has the hinder angles broadly rounded, little
produced ; the duplicature extends from the front angle almost to the other extremity,
and the lateral margin has the peculiarity of being broadest at the top, but, though
the groove is so extensive, the actual cleavage at the distal end is not very long.
The second segment has the duplicature well developed, but not externally conspicuous.
The caudal segment at the base is more than twice as broad as the total length,
which is exceeded also by the slightly convex apical margin ; the basal part of the
segment is nearh" as long as the distal, the short sides of which are only faintly
divergent.
The uropods fill the space between the caudal segment and the fifth, their outer
and distal margins forming a gentle, almost continuous, curve ; the inner rami do not
quite reach the apex of the telson ; the outer are minute, invisible from below, being
placed in a little dorsal pocket of the peduncle, at some distance from its apex.
Colour dull, mottled ; the legs and parts of the pleopods rather thickly covered
with brown markings, brighter than those on the back.
Length, 10 mm., breadth, 5 mm. The second specimen, a male, was rather smaller.
Habitat. New Britain.
The specific name is given in acknowledgment of the great assistance derived
from Budde-Luud's work on the terrestrial Isopoda.
CUBARIS ZEBRICOLOR, n. sp.
Plate LXX D.
Body moderately convex, bordered with fine hairs, appendages extremely brittle.
The pi'osepistome in frontal view convex, scarcely passing the front. The eyes not
very prominent, with about 20 ocelli. Second antennae with flagellum three-fourths
as long as ultimate joint of peduncle, this joint considerably longer than any of the
three preceding, of which the middle one is the shortest ; first joint of flagellum
half as long as second.
First segment of peraeon with the lateral edge thickened, feebly sulcate above,
with strongly marked duplicature below, the inner plate reaching as far back as the
outer, with broadly rounded apex. In the second segment the duplicature is repre-
sented by a short tooth high up.
The caudal segment is not nearly twice as broad as long, the distal part a little
longer than the broader basal part, sides strongly incurved, and then forming strongly
rounded corners to the slighth- convex apical margin.
The uropods are about equal in length and breadth, the inner rami falling a
good deal short of the apex of the telson, the outer minute, on the dorsal surface
of the peduncle adjacent to the telson just below its lateral emargination ; the apex
ON CRUSTACEA BROUOHT BY DR WILI,EY FRf>^r TFIE SOTTTIT SEAS. G57
of the peduncle of the uropods is narrow, its curve not sharply distinguished from
the line of the outer side.
Colour, forming transverse bands of dark brown, clubbed at the ends, near the
hind margin of the several segments. Tlie mesepistome is also dark, and the front
margin of the head and the first peraeon .segment.
Length of a male specimen, I'ii mm. by a breadth of 275. The specimen from
which the figures were taken was rather larger.
Habitat. Lifu, Loyalty Islands.
The specific name refers to the strikingly contrasted light and dark bands of
the dorsal colouring.
From the head to the caudal segment the dissected specimen was practically
filled with the labyrinthine parasite represented on the plate, its orange coils firmly
embracing the intestine.
AMPHIPODA.
Fam. Talitiudae.
inOO. Talitridae, Stebbing, in 'Fauna Hawaiiensis,' p. 527.
Talitrus being the earliest of the genera accepted in this family, it seems right
that the family name should be based upon it, in accordance with the example set
by Achille Costa, who in 1857 introduced the name Talitrini.
Gen. Parorcuestia, Stebbing.
LS99. Parorcliestia, Stebbing, Trans. Linn. Soc. London, vol. 7, pt. 8, p. 402.
1900. Parorcliestia, Stebbing, in ' Fauna Hawaiiensis,' p. 529.
Parorchestia h.vwaiensis (Dana).
1853. Orchestia hawaiensis, Dana, U.S. Expl. Kxp., Crustacea, p. 880, pi. 59, fig. 8,
a—g.
1899. Parorchestia hawaiensis, Stebbing, Trans. Linn. Soc. London, vol. 7, pt. >s, p.
402.
1900. Parorchestia hawaiensis, Stebbing, in 'Fauna Hawaiiensis,' p. 529, pi. 21c.
Dr Willey's specimens, which are of the female sex, were obtained at Lifu, Loyalty
Islands.
Fa.M. lillAiiDOSOMID.VE.
1895. Rhahdosomidae, Stebbing, Trans. Zool. Soc. London, v.il. 13, pt. 10, p. 366.
Gen. Itii Ai'.iiosdMA, .Vd.uns and White.
lN-i7. Rhabdosoma, Adams and White, in Wliiti's List of Crustacea in the British
Museum, p. 138.
1848. Mabdosovia, Adams and White, Zool. \ ..y. 11. M.S. Samarang, p. 63.
658 ox CEUSTACEA BROUGHT BY DE ^YILLEY FROM THE SOUTH SEAS.
1858. Macrocephalus, Bate, Ann. Nat. Hist., Ser. 3, vol. 1, p. 3(31.
1862. Rhabdosoma, Bate, Catal. Brit. Mus. Amphipoda, p. 344.
1887. Rhabdonectes, Bovallius, Bihang Yet.-Ak. Handlingar, vol. 11, No. 16, p. 39.
1888. Rhabdosoma, Stebbing, Challengei- Reports, vol. 29, Amphipoda, p. 1606.
1890. Xiphocephalus, Bovallius, The Oxycephalids, R. Soc. Sci. Upsala, pp. 3, 116, etc.
1895. Rhabdosoma, Stebbing, Trans. Zool. Soc. London, vol. 13, pt. 10, p. 367.
1900. Rhabdosoma, Chevreux, Amphipodes de THirondelle, Res. Sci. Prince de Monaco,
fasc. 16, p. 163.
Several other references, with full discussion of the genus and its name, will be
found in the writings above cited under the dates 1888, 1890, and 1895.
Rhabdosoma whitei, Bate.
1862. Rhabdosoma Whitei, Bate, Catal. Brit. Mus. Amjihipoda, p. 345, pi. 54, fig. 7.
1878. Rhabdosoma Whitei, Streets, Pr. Ac. Sci. Philad., p. 2S7, fig. 6.
1887. Rhabdosoma armatum (part), Claus, Die Platysceliden, p. 74, pi. 25, tig. 1 — 6,
pi. 26, fig. 5.
1887. Rhabdosoma Whitei, Bovallius, Bihang. Yet.-Ak. Handlingar, vol. 11, No. 16,
p. 39.
1S87. Rhabdosoma investigatoris, Giles, Journ. Asiat. Soc. Bengal, vol. 56, pt. 2,
p. 219, pi. 4.
1888. Rhabdosoma armatum, Stebbing, Challenger Reports, vol. 29, Amphipoda,
p. 1607, fig. in text of Rhabdosoma Whitei, v. Willemoes Suhm.
1890. Xiphocephalus Whitei, Bovallius, The Oxycephalids, R. Soc. Sci. Upsala,
p. 125, pi. 7, fig. 1 — 20, and numerous figures in the text.
1895. Rhabdosoma Whitei, Stebbing, Trans. Zool. Soc. London, vol. 13, pt. 10, p. 368.
Of this remarkable form two specimens, a male and a female, are included in
Dr Willey's collection. They agree well with the elaborate details supplied by
Dr Bovallius, except that the lower angle of the seventh segment of the peraeon is
less acute than in the specimens he describes. The long fourth joint of the first,
second, and third peraeopods is inflated and somewhat fusiform in the female, but
that joint in the fourth peraeopods shows no inflation. Bovallius (p. 42, 1890) observes
that m the females of the three species of this genus which he has examined the
fourth and fifth joints of the first four pairs of peraeopods ' are sometimes inflated,
and almost egg-shaped, owing to a strongly developed glandular mass surrounding
the axis of the joint for the whole of its length.' But he supposes the inflation
to be periodical, and to have some connexion with the fixation of the eggs on the
underside of the body, as he has seen full-grown females without the inflation, but
those with eggs or young ones always had it. So in the present specimen eggs
can be seen in process of development within the body. In the male specimen, the
nerve-apparatus of the eyes being broken, the prolonged ends of the pair of liver-tubes
with their large conspicuous cell.-J wear a striking appearance within the ' neck ' of the
head, as shown, though not very forcibly, in pi. 25, fig. 1, of Claus's Platysceliden.
Length of female, 55 mm., of male (with rostrum imperfect), 35 mm.
Habitat. Blanche Bay, New Britain.
ON CRUSTACEA BROUGHT BY DR WILLEY FROM THE SOUTH SEAS. 659
ENTOMOSTRACA.
BRANCHIOPODA.
Phyllocarida.
1879. Phyllocarida, Packard, American Naturalist, vol. 13, p. 128.
1883. Phyllocarida, Packard, U. S. Geol. Survey, vol. 12, p. 447.
1887. Phyllocarida, Sars, Challenger Reports, vol. 19, Phyllocarida.
Fam. Nebaliidae.
18.50. Nebaliadae, Baird, British Entomostraca, Ray Soc, p. 31.
1853. Nebaliadae, Dana, U. S. Expl. Exp., Crust., jj. 130-5.
1887. Nebaliidae, Sars, Challenger Reports, vol. 19, Phyllocarida, p. 6.
1896. Nebaliidae, Sars, Fauna Norvegiae, vol. 1, p. 6.
Dana formed the opinion that Nebalia and Chirocephalus should be regarded as
representative genera of two families in a common tribe. Claus considered that Nebalia
could best be placed in a special group Leptostraca, between the Malacostraca and
the Entomostraca.
Gen. Nebalia, Leach.
1814. Nebalia, Leach, Zool. Miscell., vol. 1, p. 99.
1896. Nebalia, Sars, Fauna Norvegiae, vol. 1, p. 7.
Nebalia pipes (O. Fabricius).
1780. Cancer bipes, O. Fabricius, Fauna Groenlandica, No. 223.
1896. Nebalia bipes, Sars, Fauna Norvegiae, vol. 1, p. 9, pi. 1, fig. 1 — 3, pi. 2, pi. 3,
pi. 4, fig. 1—8, pi. 5.
Synonymy and references to the important works on the species, genus and family
by Kniyer, Claus, Metschnikoff, Packard, and others will be found in the Fauna
Norvegiae or the Challenger Phyllocarida.
The specimens obtained by Dr Willey appear certainly to belong to this wide-
ranging species. One set were labelled " Crustacea, etc. that feed on dead fish, etc.
in the fish-traps. Lifu, Sandal Bay." Of these the largest scarcely exceeded a length
of 6 mm., and many were much smaller. Another set came from Blanche Bay, New
Britain, and were taken from (10 fathoms. Of these the largest reached a length of
7'5 mm. Baird gives a length of three-eighths of an inch for British specimens. Sars
speaks of the species reaching a length of 12 mm.
OSTRACODA.
Myodocopa.
1894. Myodocopa, G. W. Miiller, Fauna und Flora des Golfes von Neapel, Mon. 21,
p. 202.
w. V. 8'^
660 ON CRUSTACEA BROUGHT BY DR WILLEY FROM THE SOUTH SEAS.
In 1865 G. O. Sars divided the Ostracoda into four sections, Podocopa, Myodo-
copa, Cladocopa, Platycopa. In 1896 this classification is retained by Brady and
Norman. In the meantime G. W. MUUer had reduced the four sections to two tribes,
the Myodocopa and the Podocopa, the former including the Polycopidae, the single
family of the Cladocopa, and the latter including the Cytherellidae, the single family
which had constituted the Platycopa.
Fam. Asteropidae.
1896. Asteropidae, Brady and Norman, Trans. Royal Dublin Soc, Ser. 2, vol. 5, p. 628.
The genus for which this family was instituted is included by G. W. Mliller in
the Cypridinidae. But, as Brady and Norman say, " the family is widely separated
from other Myodocopa by the peculiar structure of the three pairs of maxillae, and
the presence of dorsal branchiae." They begin their account of the family with the
character, " shell more or less oblong or elliptical," which well suits some of the
species, but is not very apijlicable to the nearly circular shell of CylindroleheHs
Lobianci, Mliller. Whether the family name should be Asteropidae or Cylindroleberidae
depends on the validity or the reverse of the generic name Asterope.
Gen. Asterope, Philippi.
1840. Asterope, Philippi, Arch. Naturg., Jahrg. 6, p. 186.
1868. Cylindroleheris, Brady, Intellectual Observer, vol. 12, p. 127.
1868. Cylindroleheris, Brady, Trans. Liim. Soc, vol. 26, p. 464.
1870. Asterope, Claus, Untersuch. Grundlage der Crustaceen-systems, p. 94.
1879. Copechaete, Hesse, Ann. Sci. Nat., Ser. 6, vol. 7, p. 1.
1887. Asterope, Sars, Arch. Naturv., vol. 12, p. 188 (Ostracoda Mediterranea, p. 11).
1890. Asterope, G. W. Mliller, Zool. Jahrblicher, vol. 5, p. 238.
1894. GyUndroleberis, G. W. Muller, F. u. Fl. Golfes von Neapel, Mon. 21, p. 216.
1896. Asterope, Brady and Norman, Trans. Royal Dublin Soc, Ser. 2, vol. 5,
p. 629.
In 1840 the name Asterope was given to a genus of Echinodermata, which in
the same year received independently a different name. But whether the ostracode
was named Asterope by Philippi before the starfish was so named by Muller and
Troschel remains obscure. Since the late Mr Bladen in his Challenger report re-
linquished the claim to the title Asterope for the starfish, and Dr Brady has relinquished
his own claim to the title Cylindroleheris for the ostracode, it can scarcely be wrong
for the present to follow Brady and Norman in adopting Philippi's generic name.
Asterope, arthuri n. sp.
Plate LXXIIa.
? . Shell porcellanous, broadly oval ; antennal notch shallow, below the middle ;
rostral process broad at apex. At the opposite extremity the left valve is slightly
angled.
Vv'iLLEY. ZOOLOGICA
I.
Plate LXXI. :
ped. 3.
T.R R,S D»l.
E ffilson.Lith Caiiibridfc
Anchi c aligus nau..,jii \^/vii,icjj
ON CRUSTACEA BROUGHT BY DR WILLEY FROM THE SOUTH SEAS. 661
The eyes are oval.
The first antennae have the third and fourth joints obliquely connected, the two
together rather longer than the fifth joint, the third carrying eight marginal setae.
Of Asterope teres Norman and Brady say that the third joint is 'much shorter than
the fourth, divided diagonally so as to form two triangular joints'; but in their figure
these joints are as long as the succeeding joint, and in G. W. Mliller's figure they
are longer. In the present species the line of articulation is oblique, but does not
as in A. teres extend from base to apex of the two joints. The annulated sensory
seta of the fifth joint a little dilated at the point from which its numerous branches
spring. Sixth and seventh joints very short, the unguis long and slender, with margin
crenate or serrate.
Second antennae with eleven setae on the swimming branch, the first and last
rather short, the rest very long, four belonging to the terminal joint, the seven pre-
ceding short joints having one a-piece ; the three-jointed secondary appendage has a
single seta near the base of its third joint.
The frontal tentacle is very slender.
Mandible having the first joint of the palp produced backward at the proximal
corner in a rounded lobe armed with a plumose spine and several setae, above which
its margin is fringed with about a score of trident-headed spines; on the opposite
margin are various setae, two of them annulate and one of these very long; also at
the apex of this margin there is an unarmed laminar appendage as long as the
following joint. The tapering third joint is fringed with setae along its upper margin
and near the apex of the lower. The fourth joint has two strong annulate setae.
The first maxillae agree with G. W. MuUer's account of these appendages in the
present genus. The stem on the lower side forms a rounded projection fringed with
stout setae, and on the upper side has an unarmed tapering process adjacent to, and
more than half as long as, the gi-eat first joint of the palp, which on the upper side
is fringed above and below with setae, an intermediate space being left bare; on its
lower side it has the wonderful comb of long, parallel, terminally thickened setae, the
extremities of which form a convex line. At right angles to the apex of the first
joint is the thin second joint, followed by a minute, setiferous, apical joint.
The second maxillae consist of a slender curved plate, the concave inner margin
of v.hich is fringed closely with short setae, and having attached to it, but not quite
in the same plane, the large vibratory lamina, densely fringed on the outer convex
margin with long plumose setae.
The third maxillae (Norman and Brady) or maxillipeds or first legs (G. W. Miiller)
are in this species shaped like the blade of a battle-axe, the front margin convex,
fringed with plumose setae, and produced at each end to a somewhat acute process,
instead of having, as is more usual, one extremity rounded. In Asterope fusca, G. W.
Miiller, 1890, a small Japanese species, supposed to be devoid of eyes, the maxillipeds
are figured with both ends pointed, Miiller at that date giving this shape as generally
characteristic.
The vermiform limb has the apex beset with rows of minute teeth, and along its
distal rings there are some forty spines, some at least carrying four distal pairs of teeth.
87—2
662 ON CRUSTACEA BROUGHT BY DR WILLEY FROM THE SOUTH SEAS.
The branchial laminae are very large.
The caudal laminae (or furca) have six pairs of serrate ungues or principal
spines, the first pair of the graduated series one-fourth as long as the last. They
have above them three spinules and a row of setules.
Length, 8 mm,, breadth, 6'4 mm. Other specimens 7'5 x 5 mm., and 5 x 3"-5 mm.
Habitat. Blanche Bay, at 60 fathoms and at 70 fathoms ; the larger specimens
from the smaller depth.
The species is strongly distinguished by the character of the maxillipeds combined
with its great size. The name is chosen in compliment to Dr Arthur Willey.
Faji. Ctpridinidae.
18.50. Cypridinadae, Baird, British Entomostraca, Ray Soc, p. 176.
1880. Gypridinidae (part), Brady, Challenger Reports, vol. 1, Ostracoda, p. 151.
1887. Cypridinidae (part), Sars, Arch. Naturv., vol. 12, p. 181.
1894. Gypridinidae (part), G. W. Miiller, F. u. Fl. des Golfes von Neapel, Mon. 21,
p. 203.
1896. Cypridinidae, Brady and Norman, Trans. Royal Dublin Soc., Ser. 2, vol. .5,
p. 638.
Distinguished from the Asteropidae by the structure of the maxillae and the absence
of branchiae, or their comparatively inconsiderable development. Eumonopia, Glaus, is
distinguished as a subgenus of Cypridina, among other characters, by having a T-shaped
branchial vesicle, and Cypridina. Hilgendorjii, G. W. Mtiller, has seven branchial folds
in the male (G. W. Miiller, loc. cit., p. 20-5).
Gen. Cypeidixa, Milne-Edwards.
1838. Cypridina, Milne-Edwards, Annotated Edition of Lamarck's Anim. sans
Vertebres, vol. 5, p. 178.
1840. Cypridina, Milne-Edwards, Hist. Nat. Crust., vol. 3, p. 409.
1850. Cypridina, Baird, Brit. Entom., Ray Soc, p. 176.
18.53. Cypridina, Dana, U.S. Expl. Exp., vol. 13, pt. 2, p. 1290.
1876. Cypridina, Claus, Unters. genealogischen Grundlage der Crustaceen-systems,
'p. 92.
1880. Cyp-idina, Brady, Challenger Reports, vol. 1, Ostracoda, p. 155.
1887. Cypridina, Sars, Arch. Naturv., vol. 12, jx 206.
1890. Cypridina (and subgen.- Pyrocypris), G. W. Mtiller, Zool. Jahrbiicher, vol. 5,
p. 211.
1894 Cypridina, G. W. Mtiller, F. u. Fl. des Golfes von Neapel, Mon. 21, p. 204.
1896. Cypridina, Brady and Norman, Trans. Royal Dublin Soc, Ser. 2, vol. 5,
p. 645.
Many moi-e references may be obtained by consulting those given above. In citing
Cypridina from the edition of Lamarck's History of Invertebrates, annotated by Henri
Milne-Edwards, Brady and Norman add in parenthesis "(including subgenus Pyrocypris,
WiLLEY .Zoological R,esult
?
Plate iXXII
v.d
T R R S DtJ.
A A=;teTope rirthun n sp.
Cainbndge
B. Cypndiru). haravoni n sp
ON CRUSTACEA BROUGHT BY DR WILLEY FROM THE SOUTH SEAS. 663
W. Muller)." Milne-Edwards in 1840 declares that he only knows a single species of
the genus, C Reynaudi, from the Indian Ocean. If, therefore, that species agrees in
its characters with Pyrocypris, that name can only be a sjTionym of Cypridina, and
some other name must be chosen for any species that need to be taken out of the
latter genus.
It may be remarked that a species taken by the ' Challenger ' in the Eastern
Archipelago, and described by Dr Brady under the name ' (?) Cypridina punctata, Dana,'
in Trans. Zool. Soc. London, vol. 14, pt. 3, p. 89, pi. 16, figs. 5—9, 1897, shows a
close resemblance to a singular species from the Bay of Bengal, Eupathistoma natans,
Brady, described in the same transactions, vol. 14, pt. 8, p. 437, pi. 44, figs. 21 — 20,
1898. Probably the species ought to bear the name Eupathistoma piinctatum (Dana).
Cypridina baravoni, n. sp.
Plate LXXII b.
$. Shell smooth, oblong, length twice the height, antenual sinus below the centre,
rostral process well arched, not acute ; dorsal margin rather more convex than the
ventral, which is aj)ically produced beyond the dorsal, so as to have the ' lower half
of posterior part large gibbose' just as described and figured by Dana for his
Cypridina gibhosa (U. S. Expl. Exp., Crust., p. 129.5, pi. 91, f 4), but not so well
agreeing with Brady's Pliilumedes gihbosa (Dana) in Challenger Ostracoda, p. 160, pi.
39, f. 12. At the rostral extremity the present form agrees neither with Dana's nor
Brady's.
Eyes small, dark, rounded oval.
Fir.st antennae. First joint larger than second, third very small, fourth with a
long seta, fifth with long apical setae, perhaps belonging severally to this and two
very minute terminal joints.
Second antennae. Secondary branch apparently represented only by two or three
setae, as in the form described by Brady.
Mandibles. There are three spines or ungues at the apex ; the minute apical
joint scarcely distinguishable from the tapering penultimate joint.
The two pairs of maxillae and the maxillipeds, as shown in the figures, appear
to be consistent with the character of the genus.
The vermiform limb had but few spines on the distal rings.
The rather long narrow furca carries nine pairs of spines, the first microscopic,
and only the last three of considerable size.
Length, about 1'6 mm. Dana's gibhosa, which G. W. Muller refers to his genus
or subgenus Pyrocypris, was one-twentieth of an inch long and nearly twice as long
as high ; Brady's was of the same length, with ' height equal to more than half the
length.'
Habitat. Barawon, Blanche Bay, New Britain. Capture at night. Dana's specimen,
a female, was taken in the Pacific, lat. 15^ 20' S., long. 148° \V., Brady's, a male, in
Zebu Harbour, Philippine Islands ; both seem to be too oval in shape to be identified
with the form here described.
664 ON CRUSTACEA BROUGHT BY DR WILLEY FROM THE SOUTH SEAS.
COPEPODA.
copepoda semipaeasitica.
Fam. Lichomolgidae.
1877. Lichomolgidae, Kossmann, Zool. Ergebn. einer Reise...Rothen Meeres, Erste
Halfte, Entomostraca, pt. 1.
1880. Sapphirinidae, Brady, British Copepoda, Roy. Soc, vol. 3, p. 89.
1883. Co7-ycaeidae (part), Brady, Challenger Reports, vol. 8, Copepoda, p. 109.
1885. Sapphirinidae (Section h, Lichoviolgina), Carus, Prodr. Fauna Mediterrannae,
vol. 1, p. 3.50.
1889. Lichomolgiden, Claus, Arbeiten Zool. lust. Wien, vol. 8, pt. 3, pp. 1, 19.
1891. Lichomolgidae, Canu, Bull. Sci. France-Belgique, vol. 23, p. 477.
Concerning genera belonging or closely allied to this family there is frequent
mention or discussion in the writings of Thorell, Professor Delia Valle, Mr Thomas
Scott, F.L.S., Dr Giesbrecht, and others. Canu {loc. cit. p. 482) observes that the
members of it are doubtless near relatives of the Sapphirinidae on one side, and of
the Ergasilidae on the other. The most precise definition of it at present available
appears to be that given by Claus in 1889, and with that the new genus here assigned
to the family in many respects agrees, but whereas Claus states that the first antennae
are generally seven-jointed, they are here eight-jointed, and while he describes the
maxillipeds as three-jointed, provided in the male with powerful grappling hooks, these
parts in the new genus would rather appear to be one-jointed in the female, and to
have vanished from the male.
LiNCKIOMOLGUS, n. g.
Body pirifonn, of ten segments ; the shield, composed of head and first thoracic
segment, long and broad, the next two segments broad but short, the two following
abruptly narrower; the genital segment rather broader than long in female, but not
in male ; the four following segments very small and the caudal appendages not longer
than broad, one of their apical setae very long, especially in the female ; eyes not
perceived; first antennae eight-jointed, without ' aesthetask,' setose; second antennae
four-jointed, apically uncinate ; first four pairs of feet biramous, all the rami triarticulate,
except inner one of fourth pair, which is slender, biarticulate ; fifth feet slight, one-
jointed. Female much larger than the male, with two large egg-sacs, containing
numerous eggs.
Generic name compounded of- Linckia, name of the host, and f^oX.y6<;, skin, the
I'orm of the name being intended to link this genus with Lichomolgus historically,
even if the changes and chances of classification should sever it from the Lichomolgidae.
LiNCKIOMOLGUS CAERULEUS, n. sp., Plate LXXIVb.
Female. The shield is moderately convex, as long as the rest of the body minus
the short caudal appendages, and is considerably broader than long, with front and
sides smoothly rounded, postero-lateral angles a little produced, acute, as also are those
of the two following segments, which successively diminish in breadth, the second of
ON CRUSTACEA BROUGHT BY DR WILLEY FROM THE SOUTH SEAS. 665
them having the lateral apices obliquely truncate between two acute points, the inner
of these overlapping the two following segments. The fifth thoracic segment is longer
than the fourth, and widens distally, bearing at its lateral apices the almost rudi-
mentary fifth feet. Near each lower corner of the genital segment there stands out
a seta, not regularly tapering, but divided as it were between lash and handle.
First antennae. First two and last three joints short, the third nmch the longest,
all furnished with tapering setae, several of which are long.
Second antennae. First two joints produced each subacutely over its successor,
third carrying three small setae, apical joint forming a strong hook.
Mouth-parts. These appear to be degraded. All that I have been able to make out
clearly are first, a hook, having a broad base ornamented with very many close-set lines
running to the convex border, and a strongly curved very slender apex ; secondly, in
apparent attachment to this hook, a denticulate border lying across its base. I am
uncertain whether these parts together make up the mandible, or whether the sickle-
shaped portion is the mandible and the denticulate border the first maxilla. The second
maxillae and maxillipeds seem to be reduced to small plates with a few minute
spinules upon them. But in regard to these parts my dissections have not been
sufficiently skilful.
Feet. In the first four pairs the outer ramus has the second joint shortest, a
single spine on the outer apex of this and of the first joint, two or three on the
outer margin of the third, whicli round the other margin carries several setae, the
second joint also having one very long seta on its hind margin ; the inner ramus
in the first three pairs, though not very long, is longer than the outer, and armed
in a similar fashion, but it has a seta on the hind margin of the first joint and
no spine on its front apex, two setae instead of only one on the second joint, and
on the other hand not so large a number of setae on the third joint as in the
outer ramus ; the inner ramus of the fourth pair consists of two slender joints, the
first carrying one seta, the second with one on its inner margin, but with two setae
on its oblique apex ; the slender fifth foot is, I think, certainly one-jointed, though
the pellucid overlapping corner of the thoracic segment may give the appearance of
an additional joint at the base ; it has an oblique apex, with a rather long seta at
the outer, and a short one at the inner point. Of the caudal setae, the longest
is about equal to the body minus the shield ; on its outer side is one which seems
to vary between half and two-thirds of its length, and external to this are two short
setae ; innermost of all are one or two more of great tenuity. Length, without caudal
setae, 1"25 mm., breadth, about '85 mm. The egg-sacs contain sometimes as many as
100 eggs a-piece.
Male. Front of shield rather fiatteued, postero-lateral angles of this and the
following segment scarcely acute, and those of the second free segment rounded ;
the next segment very small, and the last scarcely defined from the gepital segment,
except by its projecting dietal angles; the genital segment with the sides parallel
instead of convex. First and second antennae and fifth feet and pei-haps the other
thoracic feet as in the female. The mouth-organs in the male seem to have become
aborted. The male and female in attachment, w'ith the shield of the male seeming
666 ON CRUSTACEA BROUGHT BY DR WILLEY FROM THE SOUTH SEAS.
to fit neatly into the strong emargination of the third thoracic segment of the female,
might easily be mistaken for a single animal, were not a warning given by the caudal
setae of the female, which jjroject from below the male, apart from his caudal
appendages. One of these couples has a rather remarkable appearance. Besides her
two packets of eggs, the female has, on one side of the genital segment, two curved,
narrow, cylindrical bodies, rather longer than that segment, possibly spermatophores.
But along with these there is another cylinder, transversely striped, as broad as the
end of the female pleon and nearly equal to the whole length of her body.
The specimens were labelled " Copepod ectoparasites from Blue Linckia. Light
blue-coloured alive."
Habitat. Feather Island, China Straits, New Guinea.
The specific name refers to the colour. The species bears in some respects a
striking resemblance to some members of the Asterocheridae, but it is without some
of the distinguishing features of that group (see Giesbrecht, Fauna und Flora des
Golfes von Neapel, Mon. 2.5), having no aesthetask on the first antennae, and, instead
of having the eggs large and few (at most 20), having them small and numerous,
reaching as above stated a hundred in number.
Panaietis, n. gen.
Head large, widening backwards, rest of the body tapering, the five leg-bearing
segments distinct, the four following separated from one another by constrictions rather
than definite boundaries ; the whole animal flat. First antennae seven-jointed. Second
antennae two-jointed, with apical hooks. Mouth-organs (seemingly) laminar. The first
four pairs of legs bii'amous, each ramus three-jointed ; the fifth pair single-jointed,
cylindric. The caudal appendages slender, about a seventh of the total length.
This genus seems to make a near approach to Anthessius, Delia Valie, and
Paranthessius, Glaus, but it is distiuguished by having the first thoracic segment
completely divided from the head-shield.
Panaietis ixcamerata, u. sp.
Plate LXX E.
Head with slightly convex front and sides, hind margin nearly straight, breadth
greater than the length. First leg-bearing segment broader but shorter than any of
the following segments ; the fifth with lateral dilatations at the middle, and the next
with such dilatations at its base ; the latter is the longest of all, the nearest to it in
length being the terminal segment, which is rather longer than the head, and has a
small apical clelt.
The first antennae are moderately stout, smooth, the second joint not thicker than
the first, but' considerably longer, the third joint short, the fourth thicker than the
fifth, but scarcely so long, about equal to the last two joints combined. In the second
antennae the lower apical hook or nail is stout, the three above it are slender, sinuous.
The appearance of the mouth is represented in the figure. It seems fitted only for
absorption of very soft or liquid food, but, being observed only in a single specimen.
ox CRUSTACEA BROUGHT BY DR WILLEY FROM THE SOUTH SEAS. 667
nothing can be said as to the variations it may undergo according to sex and stage
of life.
The first four pairs of legs arc similar, increasing in size successively. The
basal joint is large, with a fold above, the inner branch has two short stout joints
followed by a longer oval one, with three spines on the outer margin, and two or
three setules on the apex ; the outer branch is similar, but with the first and third
joints larger, the first and second each with a stout distal spine, the third with four
spines on the outer and three or four setules on the apex. The fifth legs are simple
cylinders or slender bag-like appendages attached to the prominences of their segment.
'J'he caudal appendages are rather longer than the longest segment, apparently (like
the first antennae) tipped with one or two minute hairs.
Length, 8 mm.
Habitat. Deboyne Lagoon, Panaieti, an island in the Louisiade Archipelago, New
Guinea, " from pallial chamber of gastropod." The generic name refers to the island,
the specific to the parasitic position, in which the specimen was found.
copepoda parasitica.
Fam. Caligidae.
1819. Caligidae, Leach, Diet. Sci. Nat., vol. 14, Art. Entomostraces, p. 525.
1849. Caligidae, Dana, Proc. Amer. Acad., Cambridge, Conspectus Crust., pt. 1,
p 53.
1850. Caligidae, Baird, British Entomostraca, Ray Soc, p. 25C.
1853. Caligidae, Dana, U. S. Expl. Exp., vol. 13, pt. 2, p. 1316.
1857. Caligidae, White, Popular Hist. British Crustacea, p. 310.
180 1. Caligidae, Steenstrup and Liitken, K. Danske Vid. Selsk. Skr., Ser. 5, vol. 5
(Extract, p. 9).
1864. Caligina, Krciyer, Naturhist. Tidsskrift, Ser. 3, vol. 2, p. 104.
1868. Caligina, Heller, Reise der Novara, Crust., p. IGO.
1889. Caligidae, Thomson, Trans. New Zealand Inst., vol. 22, p. 354.
1899. Caligidae, Bassett-Smith, Proc. Zool. Soc. London, p. 444.
1900. Caligidae, T. Scott, 18th Annual Rep. Fishery Scotland, part 3, p. 147.
If, in addition to the above references, the Consid. gen. sur la classe des Crustac^s
of Desmarest and Hist. Nat. des Crustaci^s of H. Jlilne-Edwards bo consulted, a tolerably
complete synonymy of the family, with the history of its extensions ami limitations, can
be obtained.
AiNcmcALKiUS, n. g.
Carapace large, scutiform. Frontal border provided with lunulae. Ventral surface
without furcula. Eyes with conspicilla. First and fourth thoracic limbs uniramous;
second and third biramous; terminal joints of first to third pairs fringed with strongly
plumose setae. Fourth thoracic segment free, small, without dorsal plates. Genital
segment without dorsal plates or processes. Abdomen small, single-jointed, the caudal
plates not fully reaching its apex.
w. V. 88
668 ON CRUSTACEA BROUGHT BY DR WILLEY FROM THE SOUTH SEAS.
This genus, though very near to Caligus, seems to be sufficiently distinguished from
it by the entire disappearance of the furcula, the peculiar character of the abdomen,
and perhaps it may be added, by the habitat of the tj'pe species.
Name from a-yx'', near, and Caligus.
Anchicaligus NAUTILI (Willey).
Plate LXXI.
1896. Caligus nautili, Willey, Quart. Journ. Micro. Sci., vol. 39, p. 145.
In general shape and proportions this species bears a close resemblance to Oaligus
abbreviatus, Kroyer, and consequently also to Caligus centrodonti, Baird, though it is
separated from both by generic characters. The carapace is only a little longer than
broad, but greatly longer than the rest of the body, in the male even more than
double as long. The genital segment is broader than long, little more than half the
width of the carapace in the female, less than half in the male. The small abdominal
segment is more than half as wide as the genital segment in the male, and less than
half in the female.
The frontal plate has a slight central notch over a small oval clear space
(described in Kroyer's species as ' orange-shaped point of attachment ') and its corners
are occupied by large almost circular lunules, abutting upon which are the first
antennae, two-jointed, the second joint much narrower than the somewhat ovoid first.
The second antennae have a hooked terminal joint, the basal seemingly longer in the
male than in the female.
Between the second antennae, a little below them, and just over the base of the
rostrum, are the two eyes, 'spectacle-eyes' according to Dana's expression (U. S. Expl.
Exp., Crust., p. 1315). Of these he says (p. 132-5), ' Besides the ordinary simple eyes
in the Caligidae, there is sometimes a pair of simple eyes with large prolate lenses
and oblate conspicilla or broad convex corneas as in our genus Specilligus.' In
describing that genus, he says (p. 1374), 'The essential point of difference between
this genus and Nogagus, is the existence of two large transpai-ent lenticular corneae
(conspicilla), exactly like those of the Sapphirinae. These conspicilla are attached to
'the exterior shell, but with some difficulty may be separated. On pressure they proved
to be brittle, though rather hard. The lenses of the eyes are situated below, near
the conspicilla, though a little nearer the median line. Between the two there is a
minute coloured spot.' The species, Specilligus curticaudis, was taken from the body
of a shark, off New Zealand. ' The pigment of the two eyes was deep blue ; the
colour of the minute spot between, bright red.' Steenstrup and Ltitken (op. cit,
p. 50) threw out the suggestion that Dana's Nogagus validus and his Pandarus brevi-
caudis, taken at the same time with his Specilligus, may be the same species. It may
be remarked that all three are clearly distinguished from Anchicaligus by the absence
of lunulae. In the species now under description the prominent eyes differ from those
described by Dana by having the conspicilla adjacent above, and the lenses a little
remote. No median spot could be perceived, but that and the eye pigment may easily
have disappeared in spirit.
ON CRUSTACEA BROUGHT BY DR WILLEY FROM THE SOUTH SEAS. 669
The rostrum apjjears to be composed as usual of slender mandibles between an
elongate upper and under lip.
The first maxillipeds have the terminal joint longer but much narrower than the
basal, with a minute spinule below the middle, and two apical setae.
The second maxillipeds have the basal joint powerful, irregularly oval, longer .than
the finger and nail ; the inner margin of the finger is rather more bulging in the male
than in the female.
The furcula is entirely wanting, and of ventral subsidiary hooks on the carapace the
only representative seems to be a spinule near the base of the second antennae. It may
be noticed that in Caligus abbreviatus Kroyer speaks of the furcula as practically rudi-
mentary. Possibly this organ is less needed in abbreviated forms.
First feet. These are slender, three-jointed, the second joint the longest, the third
short, with three plumose setae on the inner margin, the apex fringed with three little
spines and a seta.
Second feet. The stout basal joint begins with a narrow neck, perhaps representing
a true first joint, the point of junction with the true second being marked by
a plumose seta. The base carries two three-jointed rami, the first joint of the
outer and the second of the inner being the largest. The inner ramus carries one
plumose seta on the first, two on the second, six on the third joint. The outer
ramus has an apical spine to each joint, successively smaller, its second joint has two
plumose setae and the third five. The whole armature is, as so often in this pair, of
conspicuous size and beauty.
Third feet. The large and laminar, transversely oval, basal joint has the trans-
lucent wing of its outer margin divided into three portions, the middle one very small
between the two little rami. The outer ramus has a minute first joint, the second
joint with one plumose seta, the third with three such setae and some setules. The
inner ramus has a nninute first joint with one plumose seta, and a second joint with six
such setae, but two of them very small.
Fourth feet. The slender second joint is considerably longer than the rather stouter
basal joint, it is tipped with two unequal setae, and has near the middle of the
outer margin a seta, marking the point at which in some Caligidae an additional
articulation occurs. The similarity of this limb to the first maxilliped is worthy of
remark.
The linear egg-strings of the female are not so long as the body ; the eggs are
numerous, eighty or more in each string.
The caudal plates are separated by the rounded apices of the caudal segment,
which are produced a little beyond them. The minute basal joint is marked by a
setule at its junction with the obliquely oval second joint, the margin of which is
furnished with three plumose setae, successively longer inwards.
The length of the female is 4 mm., of the male, 2 5 mm.
The specimens were taken at New Britain on Nautilus ■pompilius. Others were
labelled ' from N. macromphalus. Lifu. 1896-7 etc'
Dr Willey informs me that the specimens occur " iu N. pompilius in the mantle-
cavity, and in iV. macrovi/phalus on the slimy convexity of the involuted portion of the
88— 2
670 ON CRUSTACEA BROUGHT BY DR WILLEY FROM THE SOUTH SEAS.
shell where the nuchal region of the animal plays upon the shell." In his ' Letters
from New Guinea,' Quart. Journ. Micro. Sci., vol. 39, p. 14-5, 1896, he wrote as
follows : — " Ectoparasites. In my previous note [Natural Science, vol. (5, June, 1895]
I accidentally omitted to mention the occurrence of numerous Copepod parasites in the
mantle-chamber of the Nautilus. They are present in nearly every individual that I have
examined, and are found attached to the branchiae, the internal surface of the funnel,
and in other regions of the pallial chamber. The parasites are a species of the genus
Caligus and possess the characteristic semi-lunar suckers on the first pair of antennae.
When Nautili are placed in jars the Caligids emerge in large numbers from the
mantle-chamber, and swim about actively in the water, usually in close proximity to
the sides of the vessels, ft-om which they can be removed only with some difficulty,
owing to the great adhering power of the above-mentioned suckers." A foot-note
gave the name C. nautili, pending a more detailed description. It well deserves to be
retained.
Gen. Gloiopotes, Steenstrup and Llitken.
1861. Gloiopotes, Steenstrup and Llitken, K. Danske A'id. Selsk. Skr., Scr. 5, vol. 5,
p. 36.3 (23).
1868. Gloiopotes, Heller, Crustaceen der Novara, p. 161.
1889. Lepeophtheirus, G. M. Thomson, Trans. N. Zealand Inst., vol. 22, p. 3.54.
1889. Gloiopotes, Bassett-Smith, Proc. Zool. Soc. London, p. 458.
Bassett-Smith gives the definition in the following form : — " Carapace large, oval,
scutiform. No lunulas on the frontal border. Fourth thoracic segment with two dorsal
plates partly covering the genital segment, the latter being produced backwards by two
elongated curved processes having a styliform appendage projecting from the outer border,
serrated at the edge. Abdomen long. Caudal plates lanciform. Firsfr and fourth thoracic
limbs single-branched, second and third double."
Of the two species known, the type G. hygomianus has the two dorsal plates of
the fourth thoracic segment irregularly oblong and the stiliform appendages of the
genital segment not reaching the apices of the processes from which they project,
whereas in G. huttoni (Thomson) the dorsal plates are semi-lunate, and the stiliform
appendages project beyond the apices of the prolongations of the genital segment.
The terminal caudal appendages might rather be called filiform than lanceolate ;
there is nothing leaf-like or laminar about them. The definition, as explained further
on, requires some modification now. that the male of the type species is known.
Gloiopotes hygomianus, Steenstrup and Llitken. Plate LXXIV a.
1861. Gloiopotes Hygumianiis, Steenstrup and Llitken, K. Danske Vid. Selsk. Skr.,
Ser. 5, vol. 5, p. 363 (23), pi. 5, fig. 9.
1899. Gloiopotes hygomianus, Bassett-Smith, Proc. Zool. Soc. London, p. 458.
Only the female of this species was described by Steenstrup and Llitken. The
specimen was taken in the Atlantic from some unnamed fish. Dr Willey's specimens
/ V
,/
A
r
/
\
(
►A'"
^
••/-
ON CRUSTACEA BROUGHT BY DR WILLEY FROM THE SOUTH SEAS. 671
were labelled " Rubiana, New Georgia, from tail of Albicore." The females with egg-
strings attached agree with the figures and description given by the authors above-
mentioned. The male differs from the female seemingly much more than is the case
with Mr Thomson's New Zealand species, and in a manner that will not suit the generic
definition drawn up from the female alone. The genital segment is rounded oblong, not
produced into lappets, but having lappets attached at its sides. These lappets are not
articulated but marked off by a slight constriction and extend along about two-thirds of
the abdomen or pleon ; their outer margin is folded under and ends in two spines, the
much more convex inner margin ending separately in a single spine. I am inclined to
suppose that they correspond not to the lappets in the female, but to the appendages
which project laterally from those lappets and which have the inner margin fringed with
twelve or more spinules or denticles. The dorsal plates of the fourth segment of the
trunk have theii- inner margins more divergent and oblique in the male than in the
female. In both sexes the pleon is two-jointed as described by Thomson, not unjoiuted
as described and figured by Steenstrup and Liitken. The filiform caudal appendages,
which in the female are considerably shorter than the pleon, in the male nearly equal
it in length ; at a point between a quarter and a third of their length they become
narrower and have a spinule at the outer margin ; at the apex are three unequal
spines.
Length of female, 12 mm.; of egg-strings, 12 mm.; of specimen including egg-strings,
22 mm. ; of male, 10 mm.
The agreement in habitat and general details leaves no doubt that the males
and females belong to the same species. A single specimen, like the males in size,
but with dorsal plates as in the female, had very short lappets carrying lateral ap-
pendages as large as the normal ones in the female but with only two or three
denticles as in those of the male. This may be either a monstrosity or a developing
female.
Fam. Dichelestiidae.
1898. Dichelesthiina, Bassett-Smith, Ann. Nat. Hist., Ser. 7, vol. 2, p. 91.
1899. Dichelestiidae, Bassett-Smith, Proc. Zool. Soc. London, p. 468.
1900. Dichelestiidae, T. Scott, 18th Annual Rep. Fishery Scotland, pt. 3, p. 159.
Mr Bassett-Smith in 1898 quotes from Gerstaecker a synoptic table of sixteen
genera in this family, and then modifies the table to include two new genera of his
own. In 1899 he withdraws the genus Baculus, MrAzek having shown that Bacidus
Lubbock, and Hessella Brady, are only represented by young forms of the Lernaeid
Pennella Oken. At the same time he transfers Philichthys Steenstrup, to a new family
Philichthyidae.
Bassettia, n. gen.
Head globose. Body naiTowed behind the head, with little or no demarcation of
the pedigerous segments, the genital segment oblong oval, wider and longer than all
the rest of the animal, having closely connected with it a much narrower short terminal
segment, carrying two short and narrow appendages, tipped with minute spinules. First
672 ON CRUSTACEA BROUGHT BY DR WILLEY FROM THE SOUTH SEAS.
antennae nine-jointed. Second antennae having a hooked unguis attached to a strong
basal joint. Mouth-parts e.xtremely small, apparenth^ suctorial. First three pairs of legs
unjointed, rounded, tubercular ; fourth pair at the base like the rest, but also carrying
two small narrow rami or prolongations.
This genus makes an approach to Gycnus, Milne-Edwards (Hist. Nat. Crust., vol. 3,
p. 495, 1840). That name being preoccupied must give way to Gongericola, van Beneden
(Bull. Ac. Roy. Belgique, vol. 21, pt. 2, p. .583, 18.54), which is distinguished from the
present genus by having all four pairs of legs two-branched.
Bassettia congei, n. sp.
Plate LXX F.
The first antennae have the first two and the last three joints distinct, the in-
termediate part being faintly divided into four small joints. In the second antennae
there may be a short basal joint in addition to the long one which carries the unguis.
The small oral cavity seems to contain some minute pointed organs. The tubercles
representing the second pair of legs are close to the first pair, but are larger and
more prominent. The egg-strings attached to the distal corners of the long genital
segment are somewhat longer than the animal.
Length, 3 mm.
Habitat. Blanche Bay, New Britain, parasitic on gills of Conger.
The generic name is chosen in compliment to Mr P. W. Bassett-Smith, R. N., whose
contributions to our knowledge of parasitic Copepoda are of high value.
Gen. Pseudocycnus, Heller.
1868. Pseudocycnus, Heller, Reise der Novara, Crustacea, p. 218.
189.9. Pseudocycnus, Bassett-Smith, Proc. Zool. Soc. London, p. 475.
To this genus only one species has as yet been assigned.
Pseudocycnus appendiculatus. Heller.
, 1868. Pseudocycnus appendiculatus, Heller, Reise der Novara, Crustacea, p. 218,
pi. 22, fig. 7.
1898. Pseudocycnus appendiculatus, Bassett-Smith, Ann. Nat. Hist., Ser. 7, vol. 2,
p. 368.
1899. Pseudocycnus appendiculatus, Bassett-Smith, Proc. Zool. Soc. London, p. 475.
So far as could be seen without dissection, the single specimen in the present
collection conforms with the accounts given by Heller and Bassett-Smith. The latter
writer, however, in 1899, includes in the generic account the character, "caudal ap-
pendages ver)' small, simple," whereas Heller says " cauda brevis, appendicibus valde
elongatis." The pair of long tapering appendages underneath the egg-sacs are indeed a
prominent feature of the species. Heller figures nineteen pairs of indents along the back
of the genital segment, the series stopping at some distance from its apex. In the
ON CRUSTACEA BROUGHT BY DR WILLEY FROM THE SOUTH SEAS. 673
present specimen the flattened back of the segment in question has twenty-three pairs
of indents, not quite evenly or symmetrically spaced, but covering the whole length.
The specimen is 135 mm. in length, the genital segment being about thrice as
long as the anterior part, and about twice as long as the terminal appendages. The
segment preceding the genital overlaps it with its lateral lobes and is dorsally coalesced
with it, so that it is difficult to give exact proportionate measurements. The egg-sacs
reach a length of 2.5 mm. or more.
Heller's specimens, 12 mm. long, were from the gills of an Atlantic Coryphaena;
Bassett-Smith's, 10 mm. long, were found attached to the gills of Thynnus macropterus
at Aden ; the labels with the present specimen explained it to be a Lernaeid from
the gills of Bonito, Uvea, Loyalty Islands.
THYPtOSTRACA.
(CiRRIPEDIA.)
Fam. Lepadidae.
Gen. Poecilasma, Darwin.
1851. Paecilasma, Darwin, Lepadidae, Ray Soc, p. 99.
188.S. Poecilasma, Hoek, Challenger Reports, vol. 8, Cirripedia, p. 44.
1893. Poecilasma, Aurivillius, K. Svenska Vet.-Akad. Handl, vol. 26, No. 7, p. 9.
Darwin, who rejected Trilasmis, Hinds, 1844, as an impossible name for a genus
in which the valves might be five or seven as well as only three, spells the new
name Paecilasma, but, as he derives it from 'jroiKi.Xo'i (printed Tro/ctXo?), the change
of diphthong subsequently adopted is obviously proper.
Poecilasma vagans, Aurivillius.
1892. Poecilasma vagans, Ofversigt af K. Svenska Vet.-Akad. Fiirh., No. 3, p. 123.
1893. Poecilasma vagans, Aurivillius, K. Svenska Vet.-Akad. Handl, vol. 26, No. 7,
p. 9, pi. 1, figs. 9—12; pi. 8, figs. 10, 16, 22.
Of Darwin's species only one, Poecilasma fissum, has seven valves. All the four
added to the genus by Aurivillius ia 1892 and 1893 were so provided. The species
to which Dr Willey's specimens are referable is distinguished from the others by the
circumstance that the occludent margin of the terga does not reach the orifice of
the capitulum, so that the apex of the latter is uncalcified. Also the basal edge of
the scuta is more emarginate than in other species.
Length, of the largest .specimen, 12'5 mm., by a breadth of 6'25 mm, with the
wrinkled peduncle constituting 2-o mm. of the total length, but absolutely a little
longer by reason of interlapping. Aurivillius speaks of the peduncle as § of length
of capitulum, but gives total length 12 mm., capitulum 7 mm., peduncle 4 mm., and
further says that ' the peduncle is short and thick,' and figures it in the proportion
of 2 : 5. As a matter of fact the proportions vary considerably in the spirit specimens,
but whether Aurivillius intended to convey that idea I cannot say.
674 ON CRUSTACEA BROUGHT BY DR WILLEY FROM THE SOUTH SEAS.
Habitat. Sandal Bay, Lifu, Loyalty Islands, and New Britain. Aurivillius' specimea
was found attached to the umbilicus of Nautilus umbilicatus.
Gex. Megalasma, Hoek.
1883. Megalasma, Hoek, Challenger Reports, vol. 8, Cirripedia, p. .50.
Hoek defines the genus as follows : — Valves five, approximate ; carina extending
only to the basal points of the terga, with its lower end truncated and ver}' wide.
Scuta triangular, with their umbones at a considerable distance from the rostral angle.
Mandibles with four teeth ; maxillae slightly notched, with the lower part of the edge
slightly prominent; anterior ramus of the first cirrus much thicker than the posterior
ramu.s ; the two rami of the second cirrus nearly equal ; caudal appendages uniarticulate,
short and spinose at the extremity.
Megalasma striatum, Hoek.
1883. Megalasma striatum, Hoek, Challenger Reports, vol. 8, Cii'ripedia, p. 51, pi. 2,
figs. 5—9; pL 7, figs. 8, 9.
A notable character of this species is that ' the short peduncle is quite covered
b}' the capitulum,' producing the impression at the first glance of our having to do
with a pedunculate cii-ripede devoid of a peduncle. The species is fully described
and figured by Dr Hoek. According to his description of the mouth-organs, these
show a near resemblance to those of Oxynaspis aimvillii described in this report.
Length, 9'5 mm. One of Dr Hoek's specimens was 11 mm. long.
Habitat. Blanche Bay, New Britain, on Echinus-sj)iiies measuring two to three
inches in length, and carrying also some small Balanids, Foraminifera, etc.
.J Gex. Oxyxaspis, Darwin.
1851. Oxynaspis, Darwin, Lepadidae, Ray Soc, p. 133.
1893. Oxynaspis, Aurivillius, K. Svenska Vet.-Akad. Handl., vol. 26, No. 7, p. 38.
Darwin's account of this genus is: — 'Valves 5, approximate; scuta with their
umbones in the middle of the occludent margin ; carina rectangularl}' bent, extending
up between the terga, with the basal end simply concave. Mandibles with four teeth ;
maxillae notched, with the lower part of edge nearly straight, prominent ; anterior
ramus of the second cirrus thicker than the posterior ramus ; caudal appendages,
uniarticulate, spinose.'
The single species assigned to the genus by Darwin was Oxynaspis celata, found
attached to an Antipathes, from Madeira. In 1892 Aurivillius introduced a new species,
Oxynaspis patens, also attached to an Antipathes, taken at a considerable depth off
the Island of Anguilla, in the sea of the Antilles. Besides a much larger size and
differences in the shape of the valves, this species is distinguished from Darwin's by
entii-e absence of caudal appendages, so that in regard to these the generic account
must be modified. Also to some extent the first maxillae differ in the two species.
Darwin says of the capitulum of his species, that "it seems always entirely covered
WlLLEY. ZoOLC>
A
Plate LXXIV.
Gloiopotes hvg"orma,nus.
%
pad 5
ped t
oed.2.
pe.d 3
pecC . 4.
Oxynaspis auriv:]]n. n. sp.
Koleolepas w]ile\?i- n. sp.
A a -vibaTTi. A Vv
T,R R.S. Del,
A3 vita-m. AW.
EWllrv.
Lincki oTnolgus caeruleus. n g. et sp.
Gnat, hi a aureola n. sp.
ox CRUSTACEA BROUGHT BY DR WILLEY FROM THE SOUTH SEAS. (575
by the homy muricated bark of the Antipathes, and hence externally is coloured
rich brown and covered with little horny spines. The membrane over the valves is
very thin, and is with difficulty separated from the Antipathes." But in Oxynaspis
patens the little horny spines belong to the cirripede itself, and are a kind of mimicry
of the similar spines of the Antipathes. Aurivillius is strongly inclined to believe
that in reality the same is the case with Oxynaspis celata. It is clearly true of the
new species about to bo described.
Oxynaspis aurivillii, n. sp.
Plate LXXIVc.
The capitulum and peduncle are beset, though not very closely, with little spines,
somewhat similar to those of the host. The peduncle is considerably less than half
the length of the capitulum.
The scuta are between three and four times as long as broad, widest at the
middle, the ends rounded, the upper end adjacent to the middle of the tergum, the
lower not far from the base of the carina. The terga are semi-oval, about three
times as long as broad, the convex margin adjacent to the orifice at its upper end.
The carina reaches half way between the terga, is strongly bent at the opposite
extremity, and has its apex deeply emarginate. The five valves together leave a large
part of the capitulum unoccupied.
The labrum has a convex margin, smooth in the middle, a little furred at the
sides ; the palps are rather narrow, conical, armed with several setae or slender spines.
The mandibles are setulose on surface and margins, the distal border consisting of a
moderately large separate tooth and a cutting plate divided into four smaller teeth,
of which the lowest two are nearer together than the rest, and in one mandible the
lowest has a subsidiary denticle outside. The first maxillae are setose on the outer
margin, can-y four unequal spines on the blunt outer lobe, and have the broadly conical
inner lobe fringed with sjjines of varying slenderness. The second maxillae have their
rotundo-quadrate distal margin fringed with slender spines or spinules. The cirri aro
nearly as in Darwin's tjq3ical species, with the important exception that the second
pair, like all but the first, have the rami apparently of equal thickness. Caudal
appendages seem to consist of two little, adjacent rounded plates, and therefore not
to be wanting as in 0. patens. The penis is long, and has a tuft of setae on the
narrow apex.
Colour, in spirit, pale, with brown streaks adjacent to lower part of scuta and
terga and almig the peduncle.
Length, :i mm. and sometimes a little over, the capitulum between tw-o and a
half and three times the length of the peduncle.
Habitat. New Britain, taken at 40 fathoms depth. The specimens are firmly
attached, at various angles, to the light brown scabrous branches of an Antipatharian.
The specific name is given out of respect to C. W. S. Aurivillius. By the scuta
and terga the new species is easily distinguished from the two earlier members of the
genus.
w. V. 89
676 ON CRUSTACEA BROUGHT BY DR WILLEY FROM THE SOUTH SEAS.
Gen. Conchoderma, Olfers.
1814. Conchoderma, Olfers, Mag. Gesellsch. Nat. Freunde zu Berlin, Drittes Quartel.
1851. Conchoderma, Darwin, Lepadidae, Ray Soc, p. 1.36.
1883. Conchoderma, Hoek, Challenger Reports, vol. 8, Cirripedia, p. 53.
The date and synonymy of this genus are discussed in Darwin's work.
Conchoderma hunteri, Darwin.
1851. Conchoderma Hunteri, Darwin, Lepadidae, Ray Soc, p. 153, pi. 3, fig. 3.
The specimens in the present collection agree so fully vnth Darwin's description
and figure that it is unnecessary to do more than mention the salient features of the
species. All three lobes of the scuta are narrow. The carina in two of the specimens
runs up between the terga, which are abruptly bent at the top as in Darwin's figure,
rather than in his description, for the angle formed is much, instead of little, greater
than a right angle.
Darwin gives length of the capitulum in his largest specimen as four-tenths of
an inch. The largest of Dr Willey's specimens is 21 mm. long, of which length
7'5 mm. belongs to the peduncle, the capitulum being rather over five-tenths of an
inch long.
Habitat. Blanche Bay, New Britain. Attached to cables. The specimens described
by Darwin were attached to the skin of a snake, for which the specimens here
noticed perhaps regarded submarine cables as a satisfactory equivalent. They do not
seem to have suffered from settlement on a stationary host.
Hoek says (loc. cit), " This may be a different species [from C. virgatum, Spengler] ;
but I think on account of its strong resemblance to the variety Conchoderma virgatum,
var. chelonophilus. Leach, and the complete conformity of all its interior characters to
those of Conchoderma virgatum, it would be a great deal more rational to regard it
also as a variety of Conchoderma virgatum." I have not materials for comparison, to
justify my offering an opinion on this poiat.
Gen. Scalpellum, Leach.
1817. Scalpellum, Leach, Jouru. de Physique, vol. 85, July, 1817.
1851. Scalpellum, Darwin, Lepadidae, Ray Soc, p. 215.
1883. Scalpellum, Hoek, Challenger Repoi'ts, vol. 8, Cin-ipedia, p. 59.
Scalpellum sp.
A single specimen agi-ees in general outline with Scalpellum ruhrum, Hoek, is
coloured red and white like that species, and has like it a capitulum 5 mm. long.
Also it occurs on a spine evidently of the same Echinoderm dredged in Blanche
Bay as that on the spines of which Megalasma striatum, Hoek, was found, and it
may be remarked that Hoek's two species were taken by the Challenger at one and
the same station, namely, near Luzon, in 100—115 fathoms. On the other hand the
Blanche Bay Scalpellum has a peduncle half, instead of ' about one-third,' as long as
ON CRUSTACEA BROUGHT BY DR WILLEY FROM THE SOUTH SEAS. 677
the capitulum, and the scales of the capituhim are not very prominent. Also it is
a hairy species like Scalpellum hirsutum, Hoek, with which it agrees in having the
upper latus triangular, instead of quadrangular as it is in Scaljjellum rubrinn. But
the apex is not considerably produced as in Scalpellum Mrsutum. Each of Dr Hoek's
species was represented by a single specimen, and as there is only a single specimen at
my disposal, I abstain from establishing a third of these small species, without further
opportunity for comparison and examination.
KOLEOLEPAS, n. g.
Capitulum without valves. Adhesive disk forming with the basis of attachment a
sheath for the elastic peduncle. Labrum large, with denticulate deep emargination.
Palpi strong. Mandibles with cutting edge quadripartite. First pair of cirri longer
than the rest, the rami in all six pairs shorter than the peduncle.
The same is from KoXeo?, a sheath, and Lepas, a kindred genus.
By the absence of valves this genus is associated with Anelasmu, Darwin, Alcippe,
Hancock, and Gymnolepas, Aurivillius, 1894, the last of which, having a preoccupied
title, has been re-named Erenwlepas by Weltner in 1897. Species of Alepas, Sauder
Rang, may also be entirely destitute of valves. From all of these genera the present
genus is decisively separated by the combination of characters above given. The
typical species was found in syinbiosis with a Pagurid, and the fact that the mollusc-
shell inhabited by the two in common had in some way been broken into or out of
in the immediate neighbourhood of the cirripede's position may imply that this genus
belongs to the boring groups. To these Aurivillius has lately added the family Litho-
glyptidae, with one genus and three species, which he places in the order Abdominalia,
originally founded by Darwin for the single genus and single species Cryptophialus
minutus, but subsequently augmented by inclusion of Kochlorine hamata, Noll, 1872.
H. J. Hansen in 1899 (Die Cladoceren und Cirrij^edien der Plankton-Expedition, p. 52)
considers that the group Abdominalia is untenable, as having been based by Darwin
on an entire misconception of the homologies of the cirri in the t}q3e species.
KOLEOLEPAS WILLEYI, n. sp.
Plates LXXllI and LXXIV D.
Dr Willoy's notes on the single specimen obtained supply several important details.
He describes it as a 'Sheathed and crested Cirripede living in a Turbo shell in which
was a Pagurid and on surface of which were many Actinians (seven large ones).
There was a hole in the shell exposing the end of the abdomen of the Pagurid, and
inside this hole was the cirripede attached, as shown above, to inside of shell.' The
illustration referred to (PI. LXXIV d) gives an outline of the animal with the sheath
reposing in the shallow cavity of a piece of shell, just as it came into my hands,
but the capitulum and the part of the peduncle outside of the sheath together reach a
length twice that of the sheath, without the twisting which has befallen the specimen
in spirit. Of the living form Dr Willey observes that ' It can retract itself rapidly
89—2
678 ON CRUSTACEA BROUGHT BY DR WILLEY FROM THE SOUTH SEAS.
and extend itself slowly again.' The notes on colour will be given in the course of
the general description.
The adherent disk is oval, with the narrower end towards the capitulum. On the
upper surface its skin appears to be smooth, but roughened on the lower adhesive
side. Between the two surfaces there is a pulpy mass containing numerous short
muscles. Over the peduncle, however, the sheath forms only a thin transparent skin,
and on the side next the shell this seems to be wanting or else of extreme tenuit\^
The colour of the disk in life is described as light reddish brown, a thin red line
(nigrescent in spirit) running round the translucent part which covers the base of the
peduncle, which itself is faintly roseate in life (greyish in spirit). On the under
surface, though the boundary between disk and peduncle is strongly marked, laterally
by separation and apically where they meet by colouring, yet the longitudinal muscles of
the peduncle run without flexure or any sort of interruption through the coloured band
into the disk.
The capitulum is distinguished from the peduncle by a slightly greater thickness
and by its rigidity, these characters being obviously due to its containing the chief
mass of the animal's body. The hue in life may be gathered from its description
as a white mass with a deep chocolate-bro^\^l band at its base. It is not quite
cylindrical, being laterally somewhat compressed and becoming distally carinate with a
pellucid, crest-like border, which overarches the fissure-like orifice. The sides of the
fissure close tightly together, not meeting edge to edge but with lateral compression.
They rest at the base upon a projecting bulb, and appear to be comparable wth the
corresponding part of Alcippe lampas, in which, however, there are two sharp projections
at the base, instead of a single bulb. Of this latter the function may be to give some
support to the long first cirri when protruded.
The upper lip or labrum has the free margin rather deeply concave, and fringed
with forty-six denticles. From the rounded angles two rows of fine hairs converge
backward on the surface. It has points of resemblance to the corresponding appendage
in Eremolepas pellucida (Aurivillius) and to that in Alcippe lampas, but the bullate
or swollen part extending beyond the transverse crest escaped m}' observation, perhaps
throuo-h a mishap in the dissection, rather than from the absence of a feature said
by Darwin to be common to all the Lepadidae. It is not shown in the figure of
the labrum of E. pellucida by Aurivillius.
The palps are firmly connected with the labrum, the free lobe of each projecting
in advance of or beside the rounded angle of the labrum, and having the forward
margin fringed with seta-like spines, as also the inner margin for half the depth,
behind which the lobe is emarginate. These ' palps,' though attached to the labrum,
are regarded as really palps of the mandibles. One may suppose that from the
extreme compression of the mouth-organs in the cirripedes there has resulted an
anastomosis between labrum and mandibles which has ended in the mandibular palp
havino a firmer attachment to the labrum than to its own stock. (See Darwin,
Balauidae, pp. 75, 78.)
The mandibles are rather peculiar. The upper tooth and the longer lower tooth
of the cutting edge are as usual acute, but the two intermediate processes are convex
ON CRUSTACEA BROUGHT BY DR WILLEY FROM THE SOUTH SEAS. 679
projections, the upper, which is the shorter, but the more promineut, being cut into
five denticles, the lower into ten. Between the upper tooth and the upper process
there is a close-set row of short, unequal, delicate spines, and there is similar but
less conspicuous armature at other points of the cutting edge. In one mandible the
lower tooth has a subsidiary denticle to the rear.
The first maxillae have a broad fi'ont edge, narrowing backward to a kind of
neck. The front edge has three strong spines at the upper corner, which is separated
by a notch from the rest of the margin, the spines of which are slighter, but
numerous and broken up into slightly separated groups. The cavity formed by the
neck is occupied by a thin, smooth plate. As to this the suggestion may be hazarded
that it represents the missing lower lip of the cirripedes.
Second maxillae. These appendages are represented by a pair of oblong plates
completely coalescent at the base, their outer ends rounded and fringed with spinules,
both surfaces scabrous, but the inner protruding and spinose.
Cii'ri. The first pair is the longest, and set at some distance from the rest. It is
sinuous in shape, the fii-st joint of the peduncle narrow, much longer than the second.
The rami are a little shorter than the peduncle, of five or six joints respectively, of
which the basal and the penultimate are the longest, the articulations not very distinct
and much obscured in the crowd of slender setae. The other five pairs are nearly
alike, except that the sixth appears to be rather the longest and to have a narrower
peduncle, this having in the others a very broad basal joint. The rami have seven
or occasionally eight joints, and are shorter than the peduncle. The upper ramus is
the narrower, armed only with a scanty supply of seta-like spines. The lower ramus
has here and there a slender spine, but is chiefly remarkable for the short spines on
the upper or hind margin of the last four joints, one on the short rounded
terminal joint and on the others from three to five, which are short and stout. Dr
Willey mentions that the cirri had a white spot on each.
The pleon is minute.
An immense number of small fusiform eggs were present iu the specimen.
It was taken in a fish-basket, at Sandal Bay, Lifu, Loyalty Islands.
The disk measured 1.5 mm. in length by 11 mm. in breadth. The portion of the
animal outside of the disk was about 15 mm. long, of which the capitulum occupied
<S mm., with a breadth at the crest of 5 mm. But from Dr Willey's drawing it
may be assumed that the animal can extend itself beyond the sheath to about twice the
length of the disk.
It is only fair that Dr Willey's own name should be associated with this highly
interesting discovery.
680 ON CRUSTACEA BROUGHT BY DR WILLEY FROM THE SOUTH SEAS.
DESCRIPTION OF PLATES LXIV— LXXIV.
PLATE LXIV A.
Nannastacus ossiani, n. sp. c? .
n.s. Natural size of specimen, with enlarged lateral view above, and dorsal \dew below.
a.s., a.i. First and second antennae.
m. Mandible, distal portion.
mxp. 1. First maxilliped, without respiratory apparatus (epipod and exopod).
mxp. 2. Second maxilliped.
Prp. 1, 2, 3, 5. First, second, third, and fifth peraeopods.
PLATE LXIV B.
Natmasiacus georgi, n. sp. S-
n.s. Natural size of specimen, with enlarged lateral, and less enlarged dorsal view below.
To the latter is appended a figure showing shape of the antero-lateral corner and the
pseudo-rostral projection with protruding respiratory plate of first maxilliped.
PL Dorsal view of pleon.
a.s., a.i. First and second antennae.
711. Mandible.
mxp. 2, 3. Second and third maxilhpeds.
prp. 1, 2, 4, 5. First, second, fourth, and fifth peraeopods, fifth in attachment to the
terminal segment of the peraeon.
PLATE LXIV c.
Leptochelia lifuensis, n. sp. 9 ■
n.s. Natural size of female, with dorsal view above.
oc. Eye.
a.s., a.i. First and second antennae.
m.m. Mandibles ; the right mandible above, the left below, with molar seen end on.
l.i. Lower lip.
nix. 1. First maxilla, with apex still more magnified.
ON CRUSTACEA BROUGHT BY DR WILLEY FROM THE SOUTH SEAS. 681
mxp. Maxillipeds.
ynp. 1, 2. First and second gnathopods.
prp. 2, 5. Second and fifth peraeopods.
T., urp. Telson and uropods.
PLATE LXIV D.
Leptochelia lifuensis, u. sp. S.
n.s. Natural size of a male specimen.
a.s., a.i. First and second antennae of the specimen.
gnp. 1. First gnathopod.
prp. 5. Fifth peraeopod.
PLATE LXV A.
Apanthura sandalensis, n. sp.
n.8. Line showing natural size, with hiteral view of specimen below.
c. Dorsal view of cephalon.
PI. Dorsal view of pleon.
a.s., a.i. First and second antennae.
l.s. Epistome and upper lip.
m. Mandible.
l.i., mx. 2. Lower lip and secunil maxillae in combination. The much more highly
magnified figure at the lower right-hand corner is from a second specimen.
mx. 1. First maxilla.
iii3:p. Maxillipeds.
gnp. 1, gnp. 2. First and second gnathopods.
prp. 5. Fifth peraeopod.
v/rp. Uropods in connection with pleon. Upper and lower divisions of the same shown
separately (somewhat broken).
PLATE LXV H.
Leptochelia lifuensis, n. sp. ^ .
n.s. Line representing length, natural size, with dorsal view oi aninial just below.
a.s. First antenna.
a.i. Second antenna.
l.s. Upper lip.
or. p. Ural parts below the upper lip.
gnp. 1, 2. First and second gnathopods.
prp. 1, 2, 4, 5. First, second, fourth and fifth peraeopods.
pip. One of the pleopods.
urp. U roped.
T. Telson.
682 Oy CRUSTACEA BROUGHT BY DR WILLEY FROM THE SOUTH SEAS.
PLATE LXVI A.
Gnathia aureola, n. sp.
n.s., n.s. juv. Lines indicating actual length of full-grown and younger larvae, of which
enlarged dorsal views are given above the respective lines.
L. Lateral view of full-grown larva.
Per. segm. 7. Seventh peraeon segment, embedded in the sixth.
as., a.i. First and second antennae.
ep. Supposed epistome and upper Kp.
7n. Mandible.
mx. 1, mx. 2. First and second maxiUae.
mxp. Maxilliped.
i;7ip. 1, 2. First and second gnathopods.
prp. 4. Fourth peraeopod.
pip. Pair of pleopods (the fifth).
2irp. Uropod.
T. Telson
PLATE LXVI B.
Parnnthura lijuensis, n. sp.
n.s. Line showing natural size, with lateral view of specimen below.
c. Dorsal view of cephalon, tips of tirst maxillae showing between the second antennae.
PI. Dorsal view of pleon.
n.s., a.i. First and second antennae.
l.s. Upper lip.
TO. Mandible.
l.i., mx. 2. Lower lip and second maxillae, the latter pulled aside from their natural
position facing tlie lip.
mx. 1. First maxilla.
TOxp. Maxillipeds.
gnp. 1, gnp. 2. First and second gnathopods.
prp. 5. Fifth peraeopod.
pip. 1. First pleopods.
iirj}. T. Telson and uropods in position on the pleon.
PLATE LXVII A.
Cirolana pleonastica, n. sp. (J.
n.s. Lines showing natural size, underneath the enlarged dorsal view of a specimen.
a.s., a.i. First and second antennae.
l.s. Upper lip or labrum.
m.m. Mandibles, inner surface, the left mandible on the left hand, portion of right
mandible on the right.
mx. 1, inx. 2. First and second maxillae.
ON CRUSTACEA BROUGHT I'.V DR WTLLEY FROM THE SOUTH SEAS. 683
mxp. Maxilliped, inner surface.
gnp. 1, 2. First and second gnathopods.
I'rg. 5. Fifth peraeopod.
P/p. 2, 5. Second and fifth [ileopods.
urp. Uropod.
T. Dorsal view of telson, with one uropod attached, and fourth and tiftli segments of
the ph'on.
The pleopuds, uropod, and pleon, from the specimen figured in full, the otlier detail
figures from a difl'erent specimen. The mouth-organs more highly magnified than the other
details, each set to a uniforin scale.
PLATE LXVII B.
Cirolana albicaudala, n. sp.
n.n. Lines showing natural size, underneath enlarged dorsal view of a specimen.
oc. Eye in profile view.
a.s., a.i. First and second antennae.
m.m. Mandibles, inner surface, right mandible complete, only cutting edge of left.
«ia\ 1, 7nx. 2. First and second maxillae.
mxp. Maxilliped, inner surface.
gnp. 1, 2. First and second gnathopods.
prp. 3, 5. Third and fifth peraeopods.
pip. 2, 5. Second and fifth pleopods.
T., urp. Telson (or caudal segment) and uropod.
Mouth-organs magnified to the same scale ; other details less highly magnified, but to a
uniform scale, except the lateral view of the eye.
PLATE LXVIIlA.
Hanaenolana anisopous, n. sp.
n.s. Lines showing natural size of the animal figured in dorsal view at the top of the
plate.
CD. Dorsal view of the head.
C. V. Ventral view of head (with left gnathopod attached) showing second antennae,
frontal lamina, epistome, upper lip, right mandible and part of left, spines of first maxillae,
and right ma.xilliped.
O.S., a.i. First and second antennae.
m.m. Mandibles — the right mandible on the left hnnd, the left on the right.
mx. 1, //(.'■. 2. First and second maxillae.
mxp. Maxilliped.
giip. 1, 2. First and second gnatliO])ods.
pi'p. "). Fifth peraeoi)oil.
urp. Uropod.
The mouth-organs are drawn to a uniform scale, and the more highly magnified details
also to a uniform scale ; the other appendages are less magnified than the mouth-organs,
but these likewise unitniinly.
w. V. no
684 ON CRUSTACEA BROUGHT BY DR WILLEY FROM THE SOUTH SEAS.
PLATE LXVIII B.
BenoeiJa periophthabni, n. sp.
n.s. Natural size of specimen, of which much enlarged dorsal view is given at the
top of the plate.
C.D. Head and first peraeon segment, in dorsal view, after separation from the rest of
the trunk.
C.V. The same in ventral view, one of the first gnathopods removed from the peraeon
segment to show the mouth-organs.
a.s., a.i. First and second antennae.
I.s. Upper lip.
in.m. MandiViles.
mx. 1, 2. First and second maxillae.
■mxj). MaxUlipeds.
gnp. 1, 2. First and second gnathopods.
prp. 5. Fifth peraeopod.
pip. 2. Second pleopod.
The mouth-organs are magnified to a higher scale than the other details.
PLATE LXIXa.
Cubaris cinctutus (Kinahan).
n.s. Lines indicating natural size of specimen of which a magnified dorsal view is given above.
C. V. Ventral view of anterior part of head, with first antennae and upper lip in position.
Per. s. \. Ventral view of first peraeon segment, with first gnathopod in position.
T.D. Dorsal view of pleon from the fourth segment to the telson and uropods.
T. Y. Ventral \-iew of uropods and telson.
a.s., a.i. First and second antennae to the same scale, and higher-magnification of first.
I.S., l.i. Upper lip. and lower.
m. Mandible, with distal part more magnified.
mx. 1, mx. 2. First and second maxillae partially shown, with distal extremity more magnified.
mxp. Maxilliped, with distal part more magnified.
gnp. 1, prp. b. First gnathopod and fifth peraeopod, distal part of former more magnified.
C.D. Dorsal view of head with part of first peraeon segment, from a second specimen.
In this and in the fig. T.D. the colour pattern is shown.
The two scales of magnification used for the mouth-organs are respectively higher than
those used for the other details.
PLATE LXIX B.
Philoscia lifueiisis, n. sp.
n.s. Lines showing natural size of specimen figured above.
m. Mandible.
mx. 1. First maxilla, without inner plate.
7ii.xp. Maxillipeds.
ynp. 1. First gnathopod.
prp. 4. Terminal portion of fourth peraeopod or sixth trunk-leg.
T., urp. Caudal segment with one of the uropods.
ON CRUSTACEA BROUGHT BY DR WILLEY FROM THE SOUTH SEAS. 685
PLATE LXIX c.
Paraphiloscia stenosoma, n. g. et sp.
n.s. Lines sho\\4ng natural size of specimen figured above.
oc. One of the eyes.
U.S., a.i. First and second antennae.
m. Mandible.
mx. 1, tnx. 2. First and second maxillae.
mxp. MaxUlipeds.
(jnp. 1, yiip. 2. First and second gnathopods.
prp. 5. Fifth peraeopod, or seventii trunk-leg.
pip. 2. Male organ of second pleopod.
T., urp. Caudal segment with one of the uropods, inner ramus of uropod also shown
breadthwise.
PLATE LXX.
A. Cubarin li/'uetisis, n. sp.
n.s. Length of specimen from which the figures were taken.
C.F. Frontal view of head, showing small first antenna on the right, and second
antenna (a.i.) on the left, with upper lip (l.s.) in situ below.
Fer. s. 1. V. Ventral view of part of first segment of peraeon.
T., urp. Dorsal and ventral views of caudal (telson) segment and uropods.
B. Cubans dollfusi, n. sp.
n.s. Lines showing natural size of specimen partly figured.
C.F. Frontal view of head, with upper-lip {Is.), and portion of second antenna.
Per. s. \. V. Ventral view of part of first segment of peraeon.
T., urp. Dorsal and ventral views of caudal segment and uropods.
C Cubans lundi, n. sp.
n.s. Lines showing natural size of specimen partly figured in lateral view.
CD. Dorsal view of head.
Per. s. 1. V. Ventral view of part of first segment of peraeon.
a.s., a.i. First and second antennae.
7'., urp. Dorsal and ventral views of caudal segment and uropods.
D. Cubaris zebricolor, n. sp.
n.s. Lines showing natural size of specimen from which the figures were taken.
C.F. Frontal view of head, with second antenna (a.i.) and upper lip (l.s.) in position.
Per. s. 1. V. Ventral view of part of first segment of peraeon.
T., urp. Dorsal and ventral views of caudal segment and uropods.
Par. D., Par. L. Dorsal and lateral views of parasite, with the apex more enlarged.
E. Panaielis incamerata, n. g. et sp.
n.s. Length of specimen, of which enlarged dorsal (D) and ventral (V) figures are given.
U.S., a.i. First and .second antennae.
OS. Complex of oral parts, as seen without dissection.
ped. 2. Second foot.
'JO— 2
686 ON CRUSTACEA BROUGHT BY DR WILLEY FROM THE SOUTH SEAS.
F. Bassetiia congri, ii. g. et sp.
n.s. Length of specimen, of which ventral ( V) and lateral (L) views are given.
a.s., a.i. First and second antennae.
OS. Complex of oral parts.
ped. 4. Fourth pair of legs.
c.a. Caudal appendages, attached to terminal segment.
PLATE LXXI.
Anchicaligus (n. g.) nautili (Willey).
(J , ? . Male and female in dorsal view.
n.s. Lines indicating the natural size.
S, V. Male, more highly magnified, in ^■entral view,
r. Rostrum of the female specimen.
c.p. Caudal plates of the female specimen.
L, a.s., a.i. Lunula, with first and second antennae. These and the following figures,
drawn all to one scale, from another female specimen.
■mxp. 1, 2. First and second maxOlipeds.
ped. 1, 2, 3, 4. First, second, third, and fourth thoracic feet ; of the first and second
pairs both limbs are shown with their ventral attachment.
PLATE LXXII A.
Asterope arthuri, n. sp.
n.s. Natural size of specimen, ? , of which the much enlarged lateral ^-iew is given above.
n.s*. Natural size of one of the largest specimens.
/. t. Frontal tentacle.
oc. Eye.
a.s., a.i. First and second antennae, with two spines of tirst antenna much more
highly magnified.
m. Mandibular foot.
mx. 1, 7nx. 2, mxp. First and second maxillae and ma.xillipeds.
apj). V. Appendix vermiformis, with apex much more highly magnified.
c.l. Caudal laminae.
All the details are magnified to the same scale, except the extra enlargements of parts
of the first antenna and the vermiform appendage. This last organ is from the specimen
of which the full figure is given, the other details are from another specimen.
v.d. Right valve, with body of the animal displayed by removal of the left valve.
Below are seen the branchial laminae, with the vermiform limb immediately above, followed
on the right by the eye and the second antenna, while on the left are successively tlie
maxilliped, the second and first maxillae. To the extreme left are seen protruding the ungues
of the caudal furca. Almost in the centre are the ends of the adductor muscle.
PLATE LXXII B.
Cypridina baravoni, n. sp.
n.s. Natural size of specimen, with enlarged lateral view above.
oc. Eye.
a.s., a.i. First and second antennae.
III. Mandible.
jii.i: 1, 2, mxp. First and second maxillae and maxilliped.
ON CRUSTACEA BROUGHT BY DH WILLEY FROM THE SOUTH SEAS. 687
app. V. "Vermiform appendage.
c.l. Caudal laminae.
v.cl. Dexter valve, with second maxilla in situ.
PLATE LXXIII.
Koleolepas milleyi, n. g. et sp.
Fig. at the top of plate represent.s the whole animal as seen attached to the internal
sui-face of the mollusc-shell ; or. orifice ; cr. crest.
n.s. Natural size of the animal, when dislodged; the under surface of the disk is shown;
the twisting of the peduncle exhibits the upturned orifice of the capitulum.
Cap. Fig. to the right, lateral view of upper part of capitulum ; fig. to the left, tiaee-
quarter front view of the same.
/. Labrum ; pp. palps.
m. Mandible.
mx. 1, riix. 2. One of the first and both of the second maxillae.
Cir. 1, 2, 5, 6. First, second, and sixth cirri; terminal portions from first, fifth, and
sixth more highly magnified.
T. Telson or pleon.
d. Portion of inner skin of disk.
The mouth-organs are to the same scale, more highly magnified than the cirri.
PLATE LXXIVa.
Gloioputes liygomiamis, Steenstrup and Liitken.
n.s. Natural size of the male, of which an enlarged dorsal view is given.
PLATE LXXIVb.
Linckiomolgus caeruleus, n. g. et sp.
n.s. Natural size of female, of which dorsal view in full is given in the centre, and,
to the same scale on the right, terminal part of the thorax, with male attached over the pleon.
a.s., a.i. First and second antennae. Tliese and the following details are from the female.
m., mx. 1. Mandible and fii'st maxilla, or perhaps together representing the mandible only.
ped. 1, 2, 3, 4, o. The five thoracic feet ; in ped. 4, both members of the pair are sliown.
PLATE LXXIVc.
Ox)/niispis nnrivillli, n. sp.
Specimen in lateral view much magnified.
m., mx. 1. Mandible and first maxilla.
PL.VTE LXXIV D.
Koleolepas tuilleyi, n. g. et sp.
A sketch of the animal from life, made In* l)r Willey in the East.
PLATE LXXIV E.
GiuUhia aureola, n. sp.
The animal in dorsal and lateral view, sketched from life by Dv Willey, chiclly to
indicate colour markings in the living condition.
688
ON CRUSTACEA BROUGHT BY DR WILLEY FROM THE SOUTH SEAS.
INDEX.
PLATE PAGE
Abdominalia 677
abyssorum (Anthelura) 621
aegyptiaca (Canolira) 641
Aetiobatis, parasites on 628
affinis (Calathura) 620
albicaudata (Cirolana) LXVIIb 631
Albicore, parasite on 671
AJcirona 637
Aleironidae 637
algicola (Leptochelia) 617
alloceraea (Anilocra) 640
Amphipoda 626, 657
Anceus 62-5
Anchicaligus 605, 667
angustus (Oniseus) 649
Anilocra 639
Anilocridae 639
anisopous (Hansenolana) LXVIIIa 635
Antbelura 619
Anthessius 666
Anthura 618
Anthuridae 618
Anuropus 629
Apanthma 620, 621
appendiculatus (Pseudocycnus) 672
ArmadilUdiidae 649
Armadillidium 650
Armadillo 650
arthuri (Asterope) LXXIIa 660
Asterope 660
Asteropidae 660
aureola (Gnathia) <-"._, 627
aurivillii (Oxynaspis) LXXIVc 675
Balabac, Straits of 632,638
baravoni (Cypridina) LXXIIb 663
Barawon (see also New Britain) 612,613,632
Bassettia 671
Bathynomus 629
bipes (Nebalia) 659
Blanche Bay, see New Britain
Bodotria 610
PLATE PAGE
Bonito, parasite on 673
borealis (Cirolana) 636
brachiata (Calathnra) 623
Brauchiopoda 659
brasUiensis (Leptochelia) 615
Brisson 651
caeruleus (Linckiomolgus) LXXTV b 664
Calathura 619, 624
Caligidae 667
Campylaspidae 611
Canolira 639
Caprellidae 605
Carinata (Anthura) 619
celata (Oxynaspis) 674
Ceratothoa 642
China Straits, see New Guinea
CiUoaea 643
cinctutus (Cubaris) LXIXa 651
Cinusa tetrodontis 643
Cirolana 629
Cirolanidae 628
Cirolanides 629
Cirripedia 673
Conehoderma 676
Conflict Group 633
Conger, parasite on 672
Congericola 672
congri (Bassettia) LXXp 672
Conilera 629,634
Copepoda' 664
CorUana 637
Corsica (Leptochelia) 617
crassicaudata (CUicaea) 644
Cruregens 619
Cubaris 606, 649
Cuma 610
Cumacea 606, 610
cnrticaudis (SpeciUigus) 668
cuvieri (Anilocra) 641
Cyathura 619
Cycuus, name pre-occupied 672
Cylindroleberis 660
1 For "Copepoda
' Copepoda parasitica "
semiparasitica " on
on pp. 608 and 667
pp. 607 and 664 read " Podoplea
read "Podoplea parasitica."
semiparasitica " and for
ON CRUSTACEA BROUGHT BY DR WILLEY FROM THE SOUTH SEAS.
689
PLATE PAGE
Cymodoce 644
Cymothoidae 639
Cypridina 662
Cypridinidae 602
D'Entrecasteaux Group 040
Diastylis 610
Diclielestiidae 671
dimidiata (Anilocra) 639
Diops 611
Diploexochus 649
Dolichochelia 615
doUfusi (Cubans) LXXb 654
dubia (Leptochelia) 617
duperreyi (Phyllosoma) 609
Eisothistos 619
elongata (Anthelura) 621
Entomostraca 659
Eremolepas 677
€rythraea (Paratanais) 616
Eumonopia 662
Kupathistoma 663
Eurydice 629, 634
exotica (Ligia) 040
Feather Island 600
fissum (Poeoilasma) 673
forresti (Dolichochelia) 615
forresti (Leptochelia) 615
fusca (Asterope) 601
gaudichaudii (Meiuertia) 043
georgi (Nannastacus) LXIVb 013
gibbosa (Cypridina, Philomedes,
Pyrocypris) 663
Gloiopotes 070
Gnathia 625
Gnathiidae 625
gracilis (Cirolana) 631, 632
gracilis (Philoscia) 647
Haliophasma 619
Hansenolana 629, 034
hawaiunsis (Ligia) 640
hawaiensis (I'arorchestia) 657
Hesse, Eugene .' 625
hilgendorfii (Cypridina) 602
hirsutum (SciilpcUum) 077
hirsutus (Nannastacus) 611
hunteri (Conchodenna) 670
huttoni (Gloiopotes) 070
hygomianus (Gloiopotes) LXXIVa 670
Hyssura 619
incamerata (Panaietis) LXXe 006
PLATK PAGE
indica (Eenocila) 642
inermis (Leptochelia) 615
insularis (Alcirona) 637
Isopoda 613, 045
Karuana 640
Koleolepas 605, 677
krebsii (Alcirona) 037
Lanocira 637
laticorue (Pliyllosoma) 609
latistylis (Ciiolaua) 632, 034
Leach 018, 025
Lepadidae 673
Leptanthura 619
Leptochelia 614
leptosoma (Anilocra) 640
Leptostraca 659
Lichomolgidae 064
Lifu (Loyalty Is.) 1-617,628,642,647,
1 648, 653, 654,
I 655, 657, 659,
I 669, 074, 079
lifuensis (Cubaris) LXXa 053
lifuensis (Leptochelia) LXIVc, dLXVb 010,617
lifuensis (Paranthura) LXVIb 623
lifuensis (Philoscia) LXIXb 648
Ligia 645
Ligiidae 645
Linckia, Blue, parasites on 666
Linckiomolgus 664
Lithoglyptidae 677
longirostris (Nannastacus) 611
Losilili. parasites on 640
luudi (Cubaris) LXXc 655
Ly gia 645
mariuus (Oniscus) 626
Megalasma 674
Meiuertia 642
minuta (Cirolana) 634
minuta (Leptochelia) 615
multidigita (Aega) 637, 638
multidigita (.\lcirona) 637
multidigitata (Alcirona) 637
multidigitata (Cirolana) 637
Myodocopa 669
Nannastacidae 611
Nannastacus 611
natnns (Eupathistoma) 663
nautili (Anchicaligus) LXXI 068
Nautilus macromphalus, parasite on 669
Nautilus pompilius, parasite on 669
Nautilus umbilicatus, parasite on ... 674
neapohtana (Leptochelia) 616
690
ON CRUSTACEA BROUGHT BY DR WILLEY FROM THE SOUTH SEAS.
PLATE PAGE
Nebalia 659
Nebaliidae 659
neglecta (Cirolana) 632
Nerocila 641
Nesea 644
New Britain j631, 638, 644, 647,
1 649, 656, 658, 659,
1 662, 663, 669, 672,
' 674, 675, 676
New Georgia 671
New Guinea 646, 666, 667
nigropunetata (Paranthura) 620
norvegica (Calathura) 623
oceanica (Ligia) 645
officinalis (Cubaris) 655
Oniscidae 646
Oniscoidea 645
orientalis (Cirolana) 633
OBsiani (Nanuastacus) LXIVa 612
Ostracoda 659
ovata (Eenoeila) 641
Oxynaspis 674
Panaieti 643, 667
Panaietis 605, 666
Paranthessius 666
Paranthura 619, 622
Paraphilosoia 648
Parorchestia 657
parva (Cirolana) 638
parvulus (Diops) 611
patens (Oxynaspis). 674
periophthahni (Renocila) LXVIIIb 641
Periophthalmus, parasite on 642
Philoscia 646
Phyllocarida 659
PbyUosoma 608
Pines, Isle of 617, 618, 633, 655
pleonastica (Cirolana) LXVIIa 629
Poecilasma 673
polita (Anthura) 619
Praniza 625
Procampylaspis 611
Pseudocycnus 672
PtUanthura 619
punctata (Cypridina) 663
punctatum (Eupathistoma) 663
Pyrgoniscus 651
Pyrocypris 662
quadrata (Ligia) 645
Eenoeila 640
PLATE page
Ehabdosoma 657
Ehabdosomidae 657
rhinobatis (Anceus) 628
rissoniana (Canolira) 639
rubrum (Sealpellum) 676
Sandal Bay (see also Lifu) I 616, 622, 625,
f 628, 634
sandalensis (Apanthura) LXVa 621
Saophridae 689
sarsii (Nannastacus) 611
savignj'i (Leptochelia) 617
Sealpellum 676
ScyUaiidae 609
Specilligus 668
Sphaeromidae 643
sphaeroruiformis (Cirolana) 634,636
spinosus (Diops) 611
stenosoma (Paraphiloscia) LXXXc 648
striatum (Megalasma) 674
suhmii (Nannastacus) 611
sulcata (Cirolana) 629
Sympoda 609
sjTiaca (Cubaris officinalis, var.) ... 655
Tachaea 637
Talitridae 657
Tanaidae 613
Tanais 614
tenuicaudata (Cilicaea) 644
tenuis (Paranthura) 619
tenuis (Ptilanthura) 619
teres (Asterope) 661
Tetrodon. parasites on 643
tetrodontis (Cinusa) 643
Thyrostraca 673
translucidus (Cubaris) LXTX A 651
truncata (Philoscia) 647
unguiculatus (Nannastacus) 611
uropods, Chilton, Dohrn, Gerstaecker,
on 620
Uvea (Loyalty Is.) 678
vagans (PoecUasma) 673
virgatum (Conchoderma) 676
vitiensis (Ligia) 646
weberi (Philoscia) 648
whitei (Ehabdosoma) 658
willeyi (Koleolepas) LXXIII, LXXIVd 677
zebricolor (Cubaris) LXXd 656
CAMBRIDGE : PRINTED BY J. AND C. F. CLAY, AT THE UNIVERSITY PRESS.
ZOOLOGICAL EESULTS
BASED ON MATERIAL COLLECTED IN
NEW BEITAIN, NEW GUINEA, LOYALTY ISLANDS
AND ELSEWHERE.
PART VI.
ILontion: C. J. CLAY and SONS,
CAMBRIDGE UNIVERSITY PRESS WAREHOUSE,
AVE MAEIA LANE,
AND
H. K. LEWIS,
136, GOWER STREET, W.C.
©lasgoto: 50, WELLINGTON STREET.
Jtetptig: F. A. BEOCKHAUS.
lic]s §orfe: THE MACMILLAN COMPANY.
Comtat) antr Calcutta : MACMILLAN AND CO., Ltd.
ZOOLOGICAL RESULTS
BASED ON MATERIAL FROM
NEW BEITAIN, NEW GUINEA, LOYALTY
ISLANDS AND ELSEWHERE,
COLLECTED
DURING THE YEARS 1895, 1896 AND 1897,
BY
AETHUE WILLEY, D.Sc. Lond., Hon. M.A. Cantab., F.E.S.
DIRECTOR OF THE COLOMBO MUSEUM, CEYLON.
PART VI.
{AUGUST, 1902.)
CAMBRIDGE :
AT THE UNIVERSITY PRESS.
1902 ^O
CAMBKIDGE ;
PRINTED BY J. AVD C. F. CLAY,
AT THE UNIVERSITY PRESS.
PEEFACE.
"TN bringing this publication to what I trust will prove, under the
circumstances, a fitting close, it is at once a pleasure and a duty to
express my gratitude to the various persons and bodies whose assistance
has alone rendered it possible to achieve a worthy production. It is no
part of my business to justify this method of publication of zoological
results. It is cei'tainly open to criticism and has, in fact, already received
a measure of it. It is sufficient for me to know tliat it is finished and
to record my deep appreciation of the courtesy with which the Pitt Press
have met my repeated demands up(in their consideration. The coloured
plates which illustrate my contribution to the natural history of the Pearly
Nautilus have been covered by a grant from the Publication Committee
of the Royal Society.
My work on Nautilus has been carried out in all kinds of places on
shore and aboard ship. I was not able to bring it to a conclusion before
my privileges at Cambridge lapsed and I continued it in the Biological
Laboratory at Guy's Hospital, where I I'eceived valued assistance from my
colleague, Dr T. G. Stevens, F.R.C.S.
The final touches, which always involve the heaviest labour, have been
largely added at the British (Natural History) Museum Ijy special per-
mission of the Director, Professor E. Ray Lankester.
I have also been made free of the Research Laboratory in the
Zoological Department at University College, London, by permission o\'
Professor E. A. Minchin.
Finally I am under a particular obligation to Mr A. E. Shipley in
his triple capacity of Syndic, Contributor and Friend.
ARTHUR W1LT>KY
GcLF OP Lions,
May Ut, 1902.
CONTENTS OF PAET VI.
PAGE
34. Contribution to the Natural History of the Pearly Nautilus.
By ARTHUR WILLEY, D.Sc, F.R.S.
I. Personal Narrative ........... 691
With eighteen figures. (Figures 1, .3, 4, 5, 6, 7, 8, 9, 1.5, 16 and 18 are printed
separately as 7 pages.)
Arrival at Ralum, 692; Native currency, 693; Vulcan Island, 694; Dredging in
Blanche Bay, 696 ; Commerce and Language, 697 ; Trapping Nautihis, 698 ; Ecto-
parasites, 700 ; New Hanover, 701 ; The wearing of the kabil, 702 ; Peripatus, 703 ;
The mystery of the pepe, 706 ; Jlaravot, 708 ; Styeloides evucerans, 710 ; Rhodosoma
huxleyi, 711; Spawn of Cephalopoda, 712; Malira, 713; Native fancies, 714; Tahli
Bay, 715; Tubuan and Dukduk, 716; Milne Bay and Hygeia Bay, 717; Amphisile
strigata, 718 ; Native devotion, 719 ; Ctenoplana, 720 ; Deboyne Group, 724 ; Lancelet,
725 ; Industries of Tubetube, 726 ; Ptychodera flava, 7-27 ; Prochordata, 728 ; Isle of
Pines and Mard, 730 ; Sandal Bay, Lifu, 731 ; Egg-laying of Nautilus, 732 ; Astrosclera,
733 ; Second visit to New Britain, 734 ; Appendix, 735.
II. Special Contribution 736
With Plates LXXV.— LXXXIIL, a map and fifteen test-figures.
1. Historical Survey .............. 736
2. External Form and Pigmentation 738
3. Sexual Dimorphism 740
4. Species and Range 743
5. Mantle; Shell; Nuchal Membrane 746
6. Ventral Pallial Complex 753
7. Siphuncle and Pallial Vessels 754
8. Funnel and Capito-pedal Cartilage 763
9. Digital and Ophthalmic Tentacles 767
10. Peristomial Haemocoel ; Systemic Aorta ; CephaUc Arteries 780
11. Reproductive Organs and Genital Ai'teries 784
12. Mechanism of Respiration ; Branchiae and Osphradia ; Renal and Pericardial Follicles 788
13. Eye; Rhinophore; Otocyst "93
14. The Molluscan Foot 795
15. Changes of Function, Organs and Topography 796
16. Flexure and Orientation 798
17. Morphology of the Tentacles of Nautilus 800
18. Diplomerism of Nautilus 804
19. Affinities 805
20. Food ; Migration ; Propagation 808
21. Variation and Regeneration 810
Description of Plates 813
Index °27
ERRATA.
1. "Blanche River" where it occurs should read "Blanche Bay."
2. In the article by the late Mr Bedford on Holothurians the word "topotype" was wrongly
employed at my instigation. It should read "local \-ariety."
CONTRIBUTION TO THE NATURAL HISTORY OF THE
PEARLY NAUTILUS.
By ARTHUR WILLEY, D.Sc. Lond., Hon. M.A. Cantab.
Director of the Colombo Museum, Ceylon.
1. PERSONAL NARRATIVE.
With 18 Text-figures.
LONGUM PER MARE, SED FAVENTIS UN DAE.
As I have already explained in the prefatory note which accompanied the first
Part of these Zoological Results, my journey to the Eastern Archipelago was promoted
by the Managers of the Balfour Studentship in the University of Cambridge, with the
avowed object of procuring material for the study of the embryonic development of the
pearly Nautilus. It is well known that this moUuscan type occupies a comparatively
isolated position in the series of existing animal forms, and that it is a surviving relic
of an order which was dominant in former geological epochs. It is in fact one of the
best examples of what have been called persistent types or, by way of paradox, living
fossils, that is to say, relict types of pre-tertiary creation.
It has been an object of serious investigation on the part of many of the foremost
zoologists of England, France, Germany and Holland during the nineteenth century, and
it was even one of Cuvier's regrets that he had never seen the inhabitant of the
chambered shell which had been, from time immemorial, an ornament of the conchologist's
cabinet.
Of course the chance of "acquiring a complete set of developmental stages of such
a t\^e as this, from the moment of deposition of the eggs, through the period of
incubation, culminating finally in the hatching of the miniature organism, would be
enough to whet the enthusiasm and claim the devotion of any zoologist. The dis-
tinguished German naturalist, Dr Richard Semon, whose journey to Australia, " the land
of living fossils," has resulted not only in the publication of an imposing array of
technical monographs, but also in the composition of a delightfully instructive book,
spent some time at Ambo)Tia, and tells us that a principal aim of his stay on the
Moluccas was the study of Nautilus's development'. Elated by his successes in Australia,
Semon began to imagine himself "the happy possessor of a perfect series of developing
Nautilus pompilius." In this, however, he was disappointed, and it will be seen that
my efforts were only crowned by partial success, an eventuality for which I had
prepared my mind beforehand.
1 Semon, E., In the Australian Bush, 1899, pp. 423 and 486.
w. VI. 91
692 AKRIVAL AT RALUM.
A locality where Nautilus apparently abounded had been made known, directly or
indirectly, through the instrumentality of the Wesleyan Mission in New Britain, now
known as Neu-Pommern, an island of the Bismarck Archipelago which forms part of
the German possessions ia Papua.
Thither I resorted in the autumn of 1894, generously supported by many letters
and introductions from Professor E. Ray Lankester, Professor A. A. W. Hubrecht, Sir
William Flower, Dr Anton Dohrn, the Rev. G. W. Olver of the Wesleyan Missionary
Society, Rev. W. H. Dallinger, and the authorities of the Foreign Office in Berlin and
of the German New Guinea Company.
On the way to Singapore I took occasion to call at the Stazione Zoologica at
Naples, where I had previously occupied the table of the British Association. Dr Dohrn
very kindly offered to give me an introduction to a gentleman, whose book I had just
been reading', but of whose j)i'6sent whereabouts I had no information. This was
Mr Richard Parkinson of Ralum, New Britain, whose name is well known to ethno-
graphers, and wliose house is a refuge for wayfaring strangers in those parts. The
hospitality shown to me by Mr and Mrs Parkinson, and their family, on my arrival
and during my sojourn in New Britain, was something of which I had not dreamt at
the outset of my journey, and their acquaintance with the natives, resulting from long
residence in the country, together with their readiness to do all in theii- power to help
forward my work, enabled me to commence operations without delay.
As the ship approached the anchorage opposite Herbertshohe (Kokopo) and I obtained
ray first near view of the Gazelle Peninsula of New Britain, the rising coast ^vith its
variegated sky-line, groves of waving cocoa-nut palms, scattered homesteads, with the
slumbering volcanic sentinels (the Mother and Daughters) to the right, presented
a fascinating prospect.
One of my first cares, after the first blush of arrival was over, was to procure
a boat, and this was quickly arranged by the good offices of Mr Parkiason and the
obliging kindness of the Rev. Pere Heifer, acting chief of the Catholic Mission at
Kininigunan, during the temporary absence of the Bishop, Monsignor Coupe. At this time
of the year (the middle of December) the north-west monsoon prevails, and dangerous
squalls of startling suddenness and severity are of frequent occuiTence. Two days later
the worthy Father Heifer met his doom through the swamping of a boat, in which
he was proceeding to the island of Matupi in Blanche Bay, to procure decorations for
the Christmas festivities at the Mission.
For a long time I placed much reliance upon the services of a man named
To-mangiau, who was, indeed, something of a rascal, a diable hoiteux, one leg being
shorter than the other, but not without his points and a good swimmer. Shortly after
I made his acquaintance he let himself be tattooed with a broken beer-bottle : — two
concentric circles over each breast blackened with burnt cocoa-nut. I have also seen
the natives using chips of glass and fragments of obsidian as lancets for blood-letting.
' Parkinsou, R., Im Bismarck-Archipelago, Leipzig, 1887. If I remember rightly, I owed my knowledge
of the existence of this interesting work to Dr Otto Finsch, whom I consulted at Delmenhorst before leaving
Europe.
Fig. I. Exhibition of coils of divvara at the obsequies of a native iTo-nonati
of the Gazelle Peninsula.
To /ate pagt 693.
NATIV^E CURRENCY. 693
This practice of bleeding at the seat of pain is resorted to in cases of headache,
abdominal and muscular pains.
On Christmas Eve (1894) some natives brought a white, freshly limed canoe to
Ralum for sale, the price being twenty fathoms of native shell-money or diwara',
approximately equivalent to forty shillings. As I did not possess a shell in the world
I was unable to conclude the bargain, much as I desired it. The shells (Nassa callosa,
var. camelus) are obtained by barter and by collecting from a distant locality on the
north coast of New Britain, and the bleaching, perforating^ and threading upon rattan
slips, involve so much time and labour, only good shells of even size being accepted,
that the diwara is regarded as sacred or " tambu " and is relatively as difficult to
acquire in that country as gold is in Europe. By attending a funeral on one occasion
and demanding compensation for a stolen fish-basket on another, I came into a little
propert}-, but nothing worthy of mention.
Much formality attends transactions in which " diwara " changes hands, the most
important being, of course, the purchase of a wife. The rich hoard up their wealth
in " tambu " houses, in the form of huge coils containing many hundred fathoms of
the rattan fibre with the shells threaded upon it, each coil being wrapped up in dried
banana leaves and, in this condition, resembling a life-belt. These coils are rarely
broached during a man's lifetime but, at his death, they are opened and divided into
lengths varying from less than a fathom to several fathoms, which are distributed with
singular liberality amongst those who assist at the obsequies.
The principal scene of my operations in the search for Nautilus in New Britain
was situated in Blanche Bay, a deep inlet at the north-eastern end of the Gazelle
Peninsula, which has received the picturesque native name " a bit na ta," the source
of the sea: and, indeed, it needs but little imagination, more especially in squally
weather, when clouds are gathering over the mountain-tops, to admit the poetic justice
of this ambitious appellation.
I commenced by purchasing a loosely built, palm-thatched hut on the island of
Rakai3-a or Raluan^, from a man called To-vungia*, chief of the village of Davaun, to
whom I was introduced by Mi-s Parkinson, whose knowledge of the language and usages
of these natives is very thorough. The price paid for the hut was one hundred sticks
of trade tobacco. The island itself is uninhabited, the greater part of it having been
elevated above sea-level as the result of a volcanic disturbance in the year 1878.
A small volcano, also called Rakaiya, i.e. the spirit, occurs at the base of the South
Daughter and is still simmering, sometimes emitting flames. The great drawback to
a residence on the island lies in the fact that there is no freshwater source on it.
It is now largely covered with Gasuarina trees, and near one end there is a dark
lagoon-like enclosure which is separated from the sea at low tide.
• Sometimes written " dewarra." Cf. Dr 0. Finsch, Ktluwlugische Erfahrungen aus tier Siidgee, Wien,
1893, p. 387.
^ The shells are converted into beads by dexterously knocking off the curious hump, an operation which
is performed by women.
' Also culled Vulcan Island.
* All men's names in this district begin with the prefix "To"; names of women begin with "Ya-."
91—2
694 HUT ON VULCAN ISLAND.
My hut was quite open on the lee-side, and there were rents in the thatching
which gaped wider when my boatmen abstracted handfuls of the desiccated leaflets for
the pui-pose of lighting the fire. Of course I put a stop to this practice as soon as
I detected it, and my folding bedstead and mosquito curtain, with other impedimenta,
made the place habitable for about a month, after which I moved into more commodious
quarters afforded by a Chinese trader's house at Karavia, a village lying opposite to
Raluan on the mainland.
During my tenure of the hut on Raluan, To-vungia honoured me with several
visits, generally accompanied by a crowd of retainers who clustered round the threshold
of my primitive abode, no doubt thinking that my occupation was only worthy of
a white man. When I first knew To-vungia he was unregenerate and interesting, with
the long cord-like, half-bleached, tawny, pendent, spiral locks which are characteristic
of these natives and also of some of the Solomon Islanders \
These visits were not always a source of unmixed pleasure, their inquisitive dis-
position leading some of the visitors to investigate the penetralia of my hut with
scant ceremony and somewhat to my disgust when, as too often happened, they were
afilicted with appalling sores.
Meanwhile as I had a great quantity of baggage to store away, including dredges,
laboratory appliances and other more or less necessary encumbrances, Mr Parkinson
kindly detailed some of the native labourers on the plantations, to erect a rain-proof
shelter of bamboos and plaited palm-leaves on the beach in the Ralum district at
a spot called Ka-ra-koai, meaning " under the mango-trees," of which there were three
in the vicinity. No flooring was laid down, as the black tufaceous soil is so porous
that the heaviest downpour of rain sinks immediately below the surface. Care had to
be taken not to place wooden cases on the bare earth on account of the ravages of
the white ants (termites) to which they would be exposed. Small cylinders of bamboo
(which is proof against termites) laid upon the gi-ound under the cases, afforded
sufficient protection for the time being, and I stored various articles in this place and
even worked here occasionally, although it was not possible to bring fresh material
from Blanche Bay, the distance being too great.
As I was quite ignorant of the mode of propagation of Nautilus, I employed all
available methods in my preliminary researches with the idea of aniving at my object
by a process of exclusion. These methods included the investigation of the superficial
waters of the bay by means of the tow-net, in order to determine the nature of the
Plankton or diift-fauna (Auftrieb) of this locality, since many animals which live close
in-shore or at the bottom in moderate depths (less than a hundred fathoms), lead
a roving pelagic life during the early stages of their development.
Besides pelagic larvae of Mollusca, Annelida, Echinoderma and Anthozoa, the
invertebrate Plankton also includes many adult animals which affect a pelagic environ-
ment throughout life. These are distinguished by the excessive transparency of their
tissues, which renders necessary a careful blending of the light before then- structure
' Admirable photographs of the New Britain natives taken by Mr Parkinson have been published in the
Album der Papua-Typen, edited by A. B. Meyer and R. Parkinson, Dresden, 1894.
o
696 DREDGING IN BLANCHE BAY.
can be made out under a microscope or simple lens. I have given sketches of the
Tornaria (larva of Enteropneusta), which I obtained in the New Britain Plankton in
Part III. (p. 286) of this publication. Among many other objects, a small pelagic
Mollusc known as Atlanta, belonging to the sub-order Heteropoda, was abundant at
various depths. It was my first acquaintance with this small creature which is almost
absurdly like a miniature Nautilus. It possesses in fact a perfect involute planorbiform
shell, laterally compressed, symmetrical on both sides' and not exceeding 5 millimetres
in major diameter. Of course it differs essentially from Nautilus in that the cavity
of the shell is not divided into chambers and the orifice is directed downwards,
i.e., towards the back of the animal.
Another frequent component of the Plankton consisted of the Appendiculariae,
small pelagic Ascidians which permanently retain a caudal appendage only present
during the larval phase in the life-history of the fixed Tunicata. I was somewhat
surjjrised to find specimens of a bright yellow colour and others brilliant blue, not
knowing previously that these pellucid animals ever possessed any pigmentation. It
has, however, recently been noted by Mr E. T. Browne that specimens of the genus
Oikopleura occurring in British waters are sometimes coloured a bright crimson^.
More important than these Plankton excursions, which generally took place in the
small hours of the morning before sunrise, was the investigation of the nature of
the ground at the bottom of the sea in Blanche Bay. This revealed the existence
of submarine chasms probably inaccessible to the dredge, and it may be at once noted
as a possibility that such protected situations are naturally chosen by Nautilus for the
purposes of nidification and oviposition.
In order to clinch matters at the outset I seized the opportunity of engaging,
by favour of Herr Thiel, the representative of the firm of Messrs Hernsheim and
Company on the island of Matupi in Blanche Bay, the services of a small steam-
launch, the only one in the Archipelago. With the kind permission of Mr Adam
Sedgwick I had taken out with me from Cambridge a couple of drums of wire-rope,
and one of these was fixed upon thwarts on the launch. Eventually I came to the
conclusion that, when dredging from light craft in moderate depths, ordinary strong
hemp rope is to be preferred. A pair of gun-metal bollards were screwed on to the
stem of the vessel for the rope to pass over when hauling in, since, without these,
the wire would cut through Australian hard wood with the greatest ease. The depth
varied from thirty to seventy fathoms and there is no doubt that the result of these
laborious dredgings was not very encouraging. They simply proved (apart from the
chasms mentioned above) that the ground on which Nautihis is caught in the fish-baskets
is essentially composed of volcanic mud and pumice stones.
While staying at various places about Blanclie Bay my food, besides tinned stuffs
and an occasional fish, consisted principally of yams in the north-west season and taro
during the south-east monsoon. These invaluable tubers and corms are purchased from
the women at the markets which they hold periodically on the beach. They bring
' Not in all species, of. Soiileyct's figures in the Atlas of Voyage de la Bonite, 1836 — 1837.
- "The Fauna and Flora of Valencia Harbour," P. Irish. Ac. (3), v. p. 751, 1900.
Fig. 3. Market-women at Ralum protecting themselves with pandanus rain-mats during a shower.
I'o face im'.K 697.
COMMERCE AND LANGUAGE. 697
dovm astounding loads of vegetables from their gardens in the bush, carrying them
on their backs in palm-leaf baskets, the weight being supported by a band which
passes across the forehead. Besides this vegetable load they frequently carry an inflant
as well, slung in front in a native bark-cloth fastened over the shoulders. All the
infants at a certain age have their small bodies beset with numerous raised sores
nearly an inch in diameter. To the inexperienced ej-e they present a shocking ap-
pearance at this stage, but one soon learns that the mothers cherish these sores and
even, I believe, keep them open. If the sores do not break out there is cause for
anxiety for the future health of the child. The markets are aboriginal institutions
and are held in the bush, far from the influence of the white man, as well as on the
beach.
As a general rule fish caught in the fish-baskets can only be bought with diwara,
while yams and tare may be paid for with tobacco.
The natives are bom tradere in their owti way, and liberality i.s likely to have
a demoralising effect upon them. But their ways are peculiar, and appear at times
unreasonable ; in fiict their actions are often quite devoid of reason, being based either
upon tradition or inherited instinct or else upon some chronic, deep-seated, primary
logical fallacy to which they are held in ineradicable subjection. Unfortunately besides
these primal, more or less picturesque attributes, they well understand the subtleties
of double-dealing and sophistry. If, for example, one applies soothing balsam to their
sores, they are ready to assume that the obligation rests with the donor, unless perhaps
their idea is that a little tobacco is necessary to complete the cure. On one occasion
the paddles had been left for a few moments unguarded in my canoe on the beach.
A passing native promptly appropriated them, took them to a neighbouring compound
and had the impudence to ask for payment in response to my indignant demand for
restitution.
It is not difficult to acquii-e a cursory knowledge of the New Britain language,
which possesses simple detached particles and a straightforward sjTitax'. Of course
the natives spend a good deal of their time lounging about or idly amusing them-
selves, and the word they use to express this pastime is " limlibur." Often during my
operations in the Bay or on shore, natives whom we passed in canoes or on the
beach inquired whether I was amusing myself in the sense of "limlibur."
There is no telling for how many years the Nautilus-&shery has been carried on
by these natives. It is not a regular source of food but is resorted to when other
provender is scarce. The range in size and age of the individuals taken in the fish-
traps is considerable, and as the young are tender and tempting, it happened that
I had some difficulty in impressing upon the fishermen that it was precisely these
which I jjarticularly desired to procure. They had no knowledge whatever of the
possible egg-laying properties of tiie " pal-a-lialia'^" on the contrary they scouted the
' An exceedingly useful and reliable dictionary, following the plan adopted by the Kev. G. Brown for the
language of the Duke of York Island, is entitled " A Dictionary of the New Britain Dialect •««♦•••••* also
a Grammar" by the Rev. R. H. Rickards, 1889. It has not been printed, but I am informed that there is
a copy in the library of the British Museum.
- The native name for Nautilus.
698 TRAPPING NAUTILUS.
idea with ridicule. This was in itself a bad sign, in consideration of the intimate
and intuitive acquaintance which they manifest with many natural phenomena.
In New Britain the year is divided into two sharply contrasted seasons distinguished
by the direction of the prevailing trade-wind. The fine south-east season lasts approxi-
mately from April to October and during the rest of the year, from November to March,
the north-west wind blows almost unceasingly, bringing in its wake rain and squalls.
In other localities, such as the Moluccas and the Philippines \ Nautilus can only
be obtained during the time of the south-east monsoon, but Blanche Bay has this
advantage, that it yields up its treasure throughout the year.
The traps which are set for Nautihis at a depth of from thirty to seventy
fathoms are baited with small soft-skinned fishes which congregate in shoals at various
points of the coast and are obtained by exploding a charge of djmamite in their midst.
This mil no doubt appear a reprehensible method to an angler but it should be
remembered that it is the common custom, sanctioned bj- high and low in these parts.
On a typical occasion in January 1895, which I may describe, having procured my
bait I took it to Davaun, To-vungia's \411age, in order to make arrangements for the
setting of the traps during the coming night. I also purchased for myself a fish-
basket in exchange for a " lava-lava " (waist-cloth) and six sticks of tobacco, and then
returned to Vulcan Island. As soon as the fishermen who happened to be there saw
that I was becoming independent of them they set to work with a will to bait the
trap by tjing the small fish called " malabur " on fibres purposely suspended inside
from the framework. After this had been done and a heavy stone had been attached
at each end the fish-trap was ready to be mounted upon a canoe and taken to the
selected spot.
We baited two traps on this occasion and started out, in two canoes, shortly
after sundown, paddling towards Davaun and stopping rather close to the shore
opposite to a point where a light was burning at " house belong To-galli," all positions
being ascertained by dead reckoning. We began to lower my basket at 7.20 p.m.,
using a native rattan cable, and finished paying out the rope after the lapse of
twenty minutes; during this time, that is to say while the basket was slowly sinking,
the canoe was kept in gentle and silent motion by the assistants.
To-kiap, master of the canoe, having made a float of light wood, set the whole
thing adrift and then we cruised about until 10 o'clock when we recovered the float
and began to haul up. There were no Nautili in the basket, a disappointment of
small moment to the biologist accustomed to negative results.
A thermometer, kindly lent by Mr Parkinson, which I had taken the opportunity
of fixing to my basket, showed a difference of about 9^^ C. between the temperature
of the bottom of the bay and that of the top, the respective readings being 20° C.
on the i^awttiiis-frequented ground as against 29° C. at the surface.
1 Coucerning the occurrence of Nautilus at Amboyna, Dr Semon's work. In the Australian Bush, may be
consulted. As for Negros, one of the Philippine islands, I am indebted for information to Prof. D. C. Worcester,
formerly of the University of Michigan, and to Prof. Bashford Dean of the Columbia University, Xew Tork.
Prof. Dean has published an illustrated account of his observations, entitled " Notes on Living Nautilus," in
the American Naturalist, Vol. xxxv. 1901, pp. 81'J— 837.
fcf
'A,
^■^
a^:
^m^
Fig. 4. Natives baiting a trap for Nautilus on Vulcan Island,
Blanche Bay.
Fig. 5. Lifuan fish-trap on raft with native (Saonoi; the raft is moored upon
the beach in Sandal Bay.
[Phatogrnph by Rn: J. Hmlfidil.\
To /ace page 699.
TRAPPINC4 NAUTILUS. 699
We then moved on a short distance until we an-ived opposite " house belong To-
gogi " ; here we lowered once more, hauling up again shortly after midnight. This
time we were fortunate in the capture of two specimens, a large mature male and
an immature female. The second canoe now rejoined us in order to hand over their
bag which amounted to six specimens, and we then returned to Vulcan Island. On the
following morning between 7 and 8 o'clock, To-vungia came across from Davaun
liringing seven more Nautili caught overnight.
Of the fifteen individuals procured on this occasion only two were females and
this numerical disproportion between the sexes was maintained, with greater or less
variation, in the subsequent takings.
One more incident may suffice to complete my description of the Nmdilus-^shery
in Blanche Bay. On January 21, four canoes, each can-ying a baited fish-trap, left
Vulcan Island at 5.30 p.m. to sink the baskets on the Nautilns-gvownA.
I accompanied them as before in To-kiap's canoe. Having cast off the floats we
went ashore at Davaun to rest, lying down on plaited cocoa-nut leaves placed on the
gi'ound. At about 10 o'clock, a gale of wind and rain burst upon us and I adjourned,
with To-mangiau and To-kiap, into the latter's house, a good weather-proof palm-leaf
hut, but veiy small. There were already two men asleep on the ground and a fire
burning in the middle. However, we went in and lay down to sleep through the
gale which lasted until midnight, and it was not before 1.30 a.m. that our members
were sufficiently roused to be able to set out for the pui-pose of raising our baskets. My
basket contained six Nautili, and altogether the catch amounted to twenty-one, of which
sixteen were males. Upon commencing to return to Vulcan Island, the clouds looked
so black and threatening ahead, lowering ominously over the summits of the Mother
and Daughters, that we reluctantly deemed it necessary to put back to Davaun.
Accordingly we returned to enjoy the shelter of To-kiap's roof once more and hardly
had we regained it when the storm-clouds broke and converted the bay, for the time
being, into a howling wilderness. We had in fact just been able to haul in our traps
in the interval between two severe squalls.
Just as the local inhabitants regard Blanche Bay as the source of the sea, so as
I lay prone upon my mat listening to the raging of the elements it seemed to my
fancy to be the veiy cauldron in which the north-west squalls were brewed.
Nautilus is thus trapped in the watches of the night and it is " tambu " to speak
while the lines are being hauled \\\^ from the depths. At first I was liable to
desecrate the performance by breaking the silence to utter some jargon of " pidgin
English " or perhaps mutilated patois, but I soon learned to respect the unwritten law.
The fish-baskets are large barrel-shaped contrivances made by interlacing bamboo-
.slips bound together by rattan fibre. They are five or six feet in length and three or
four feet across the middle ; they are furnished at each end with an inverted funnel
terminating in a forest of loose slips which lie across the inner mouth of the funnel
and, while not preventing the entrance of various marine animals such as Nautilus,
fishes, sea-urchins and Crustacea, effectually foil their attempts to escape.
The construction of these fish-traps varies in certain particulars of minor import-
ance although carefullv considered by their makers. When not in use they are stored
w.vi. " " 92
700 ECTOPARASITES OF NAUTILUS.
away by being suspended from the overhanging stems of the cocoa-nut pahiis which
fi-inge the beach, and consequently they are a conspicuous feature in the shore scenery.
The best of them are employed for surface-fishing during the south-east season ; for
this purpose they ai-e attached to large bamboo floats which are held in position by
anchors made by enclosing heavy stones in a rattan cage. When once they are placed
in position these anchors are never moved and frequently my di-edges brought up lengths
of former rattan cables which had become water-logged. In the course of time these
anchor ropes become coated with Hydroids and also serve as a resting-place for Annelid
worms (especially one handsome species called Amphinome rostrata). Sometimes gi-ape-like
bunches of Cephalopod eggs are found attached to them. Thinking that Nautilus might
use them for this purpose I once requested the Rev. W. J. Chambers of the Wesleyan
Mission to use his influence to induce certain natives to allow me to haul up two or
three of theii- anchors, but the result was negative.
The opening of the fishing season, when the natives go out in their white canoes to
lay their anchors and set their traps, is a joyous occasion and is heralded by the blo^ving
of Tiiton conchs.
The squid egg-clusters referred to above are called "a mo a tauka" and I made
special inquiries of the natives to ascertain whether they were acquainted with " a mo
a pal-a-lialia." They replied No with emphasis ("pata-na") as if inci-edulous of their
existence. This was sufficient to convince me that the eggs of Nautilus are not deposited
in obvious bunches or in ordinary situations.
When living Nautili are confined in a vessel of water, numbers of small Crustacea
issue from the mantle cavaty in which the gills lie and also from the funnel by the
pumping action of which the shell is driven backwards when the animal is swimming
freely. These ectoparasites apparently live upon the slime which is secreted from the
mucous surface of the body, and, on leaving thefr host, swim about actively in the
water and also cling tenaciously by means of a peculiar suctorial apparatus to the sides
of the vessel'.
In the month of February (189.5) I gladly accepted an invitation from Mr Parkinson
to accompany him on a short recruiting voyage to New Ireland and New Hanover in a
small 50-ton fore-and-aft schooner called the Maj^flower.
There was the possibility that a locality might be found where Nautilus could be
investigated in shallower water than in Blanche Bay. But these countries are too ^vild
and no reliable information could be obtained. The journey was, however, interesting
in itself as providing a glimpse of native systems with quite a different code from
that by which the inhabitants of New Britain are bound, notwithstanding the fact that
the islands form part of one and the same compact archipelago. The distances are too
great for small canoes to traverse so that there is practically no regular communi-
cation between New Britain and the larger islands except the Duke of York Island.
Moreover the natives are suspicious not only of the white man but of their own colour.
Between New Ireland and New Hanover there is a labyrinth of islands and
' They are named Anchicaliniu^ nautili by Mr Stebbing, Part v. pp. 667 and 668. For an account of
Crustacean ectoparasites of other Cephalopods see Wierzejski, k., " Ueber Schmarotzerkrebse von Cephalopoden,"
Zeitschr. wiss. Zool, Bd. 29, 1877, p. 562.
NEW HANOVER. 701
channels, one of the islands when seen from the west presenting a symmetrical appear-
ance with a flat-topped mountain in the centre, which has suggestefl the appropriate
name of Mausoleum Island.
The opening at the top of the New Hanover and Mausoleum Island canoes was
very narrow, the cavity of the canoe widening out inside ; the outrigger was adorned
^vith pigs' tails. Some of the natives had a wild hunted expression in theii' eyes
such as I had not seen elsewhere ; one fellow was decorated with a lurid red pigment
on chest, shoulders, back and thighs. Whether this was for mere display or to produce
a terrifying effect, I do not know. On the upright pole of the outriggers there was
occasionally a Nautilus shell, no doubt placed there as an object of beauty, the natives
being very susceptible to singularity of form, although the shell is also used for the
more commonplace function of baling out the canoes. Many of the canoes only came
within shouting distance, their crews being doubtless smitten by their consciences on
account of former delinquencies and afraid to come alongside. Higher up along the
coast of New Hanover we went ashore and engaged the bulk of the boys, the natives
being used to the recruiting business.
The parts visited on this occasion included the whole south-west of New Hanover
and along the north coast round Cape Charlotte as far as the island of Kung, one of
the North Islands, where there was a trader's station. On the return journey we passed
through Steffen Strait as far as the island of Nusa and Cape Nowon on the main-
land of New Ireland.
The men of New Hanover go about in a state of nature, carrying wooden spears either
singly or in sheaves. Some of them wear armlets of closely woven fibre which sometimes
become so tight, with increasing age, as to be almost concealed below the neighbouring folds
of flesh. They also wear spindle-shaped, well-fashioned rods of shell passed through the
nasal septum, just as in New Britain some of the better class of bushmen employ quills from
the cassowary {Gusuarius hennettii) as nose-sticks. Occasionally one sees a man wearing
a large white disc cut out of the giant clam Tridacna and suspended by a string round
the neck, forming a kind of medallion varying from two or three to five or six inches in
diameter. This ornament (called "kapkap") is more frequently met with in New Ireland
and is generally much improved by the addition of a circular lamina of tortoise-shell
beautifully fi-etted, the whole forming an artistic composition, the tortoise-shell plate being
thrown into elegant relief by the white background furnished by the Tridacna shell-disc.
Armlets made by cutting out segments from large froc/tzts-shells are counuonly
worn by well-to-do women throughout the Bismarck Archipelago. But the woven
armlets mentioned above serve a double purpose, decorative and utilitarian, since all
kinds of things, such as a pipe, ;i stick of tobacco, ornamental leaves and so forth can
be carried thrust in between the ring and the flesh.
The women of New Hanover and neighbouring islands wear very curious liats
made of pandanus leaves, called " kabil," which they are unwilling to remove in the
presence of men. When I first saw women i-unning along the shore to meet the boat,
I was greatly fascinated by this singular head-gear which somewhat resembles a bishop's
mitre. The scenery in New Hanover is very attractive and is not rendered less so by
the native taro plantations in the form of terraces on the sides of the well-wooded hills,
92—2
70:3 THE WEARING OF THE KABIL.
with palms and Cycads in the tbregronnd. They use long, light: nairow rafts made with
the midribs of sago-palm leaves lashed together and propelled with simple bamboo rods.
When a crowd of natives came to the beach the women kept together in a separate
gi'oiip, and when a young fellow accepted the recruiting terms and was taken away in the
boat the poor women often raised a terrible wail, shrieking, stamping and rubbing sand
over their bodies. But from previous experiences I felt that such gi-ief would be so<in
assuaged, heart-rending though it appeared.
The wearing of the "kabil" is an act of deference to the men. In one village called
Neila-as, on the island of Kung, there was a tree near the chief's house, upon which
several skulls were exposed, with huge rents behind the ear. There can be no doubt
that in this country infringements of the social code are visited with swift retribution.
At Neila-as we wanted to photogi-aph a group of the women •w'ith their kabils,
and in order to place them in a favourable light it was necessary for them to cross
the village-square. I was much interested to observe that they crossed this space on
their knees in view of a group of men who were watching them.
Birds are " tambu " or sacred in New Hanover and each indi\'idiial native acknow-
ledges a certain kind of bird as his patron-bird and the people range themselves
accordingly into groups or phratries, named after the birds. The esteem in which birds
are held is further manifested in their car\'ings, particularly in the figure-heads of
their canoes which are carved out of the same piece of wood fi-om which the dug-out
itself is made ; the head and eyes are usually distinct but the rest of the body is
drawn out into a fantastic scroll and in fact the whole figure of the bird may
degenerate into a decorative scroll. Another kind of carvings representing birds is called
"kui." This is used in dances and is provided wth a spatula-shaped basal piece by
which it is held in the mouth ; the eyes are the blue opercula of Turbo petholatus.
Seismic disturbances are of frequent occurrence in New Britain and when they
occur at night the bush re-echoes with the blowing of conchs and the tattoo of
" garamats " or native tomtoms, to soothe the angry spirit. If there were any houses
built of stone the earthquakes would no doubt frequently be classed as disasters, but
under present conditions only minor casualties occur such as the upsetting of lamps and
other household implements.
In the Strait of Rakaiya between Vulcan Island and the mainland, both men and
women used to come do^vn to scoop up the black sand or mud near the shore with
small hand-nets in search of the " diwara " shell which occurs here in small quantities but
is said to be of inferior quality', and varies in colour from almost pure white to jet-black.
The natives sift the sand through their nets and each shell, as they fijid it, is carefull}'
stored away in their mouths until their jowls are full.
Finding it necessary to have some lighter craft than a whale-boat for general
purposes, and not being able at that time, for reasons stated above, to purchase a New
Britain canoe, I commissioned a Samoan mission-teacher to make me a canoe in Samoan
fashion during his spare time. In this plain unadorned dug-out I was able to traverse
the whole extent of Blanche Bay with a single companion. When it sprung a leak, as
all canoes do sooner or later, the holes were stopped up with the bruised fruit of a tree
' The principal source of the shell lies far away on the north coast in the district of Beining.
-?'
Fig. 6. Bird-scrolls ("kui"), New Hanover.
Fig. 7. Domestic scene at the village of Neila-as. New Hanover.
T»/»Cf ]HHIr 702.
PERIPATUS IX NEW BRITAIN. 703
called " a tita" (Parinanum laurinum) which has a wide distribution and is even called
by the same native name in the Solomon Islands, where it is employed for the same
purpose and also as cement for the remarkable inlaid work which the Solomon Islandei-s
practise in the decoration of their canoes, paddles and weapons.
The kernel of the nut is removed and rubbed against a stone to produce a pulp.
According to native directions " u na ud ia ta ra wat," which means, word for word,
■' you must rub it on the stone," the particle " na "' implying both obligation and futurit}-.
A quantity of dust or ashes is mixed with the pulp to form a paste which is then
employed like putty in filling u]j the crevices of the canoe. The composition soon turns
brown and dries hard, after which it is plastered over ^vith lime obtained from calcined
corals and mixed with the expressed juice of the cocoa-nut.
The natives are cynojjhagists and I was told that they rub the bruised fruit
of the Barringtonia speciosa called " a vutun " on the closed eyes of recently born
pups, to cause them to open, but I did not actually see this veterinary feat performed.
I stayed at Karavia, renting about half a bungalow belonging to Ah Tarn of
Matupi, from March 1 to July 12 (189.5). During this time hardly a day passed
Nvithout some expeiiments, operations or observations relating to Nautilus being carried on.
If I made an excursion into the bush behind Karavia to shoot a pigeon for lunch
or for change of occupation, either a cage, barrel, basket or hempen tangle would be
testing its quality in the silent depths of the bay.
It was during one of these excursions, on the occasion of my second visit to
New Britain, that I had the good fortune to come across a species of Peripatus.
At the outset of my journey I had calculated upon the possibility of discovering the
Peripatus of New Britain, but it was not until near the close of my voyages, in the
year 1897, that my dream was realised.
It is well known that Peripatios occupies an isolated position in the animal kingdom,
presenting signs of affinity both to the Annelid Worms and to the Tracheate Arthropods
but not falling into the sequence of normal gradation of forms. When seen from above,
immediately after turning over a stone or a fallen leaf or a piece of rotten bark, it
resembles a catei-pillar ; if looked at from below it is like a centipede with many pairs
of clawed legs, and when held in the hand or between the fingers it is found to be
soft like a caterpillar or a worm.
From a zoogeographical point of view, Peripatus possesses a perennial interest which
does not diminish as time goes on, although the recent discovery of a new genus, with
three new species, in the Malay Peninsula during the Skeat Exjiedition to this region,
renders generalisation for the present impossible. We may confidently await further
jidditions to our knowledge of the distribution of the Onychophora for many yeai-s to
come, according as the various islands of the Eastern Archipelago are searched for their
cryptozoic inhabitants. For example it may seem difficult to believe that Peripatus has
been overlooked in Java which has been visited by so many naturalists of the highest
distinction, but it is equally incredible that it does not occur there. It is to be looked
for under leaves, bark and stones in damp mountain valleys near streams or springs, and
its absence from the neighbourhood of one source does not preclude the possibility of its
occurrence near another half-a-mile distant. It is essentially a creature of contradictions
704 PERIPATUS.
both in its structure and habits. While abundant moistui'e is absolutely indispensable
to its existence, immersion is quickly fatal ; it is therefore likely to prove of peculiar
theoretical value to the zoogeographer since the only conceivable means of transit from
one place to another is by land. But the occurrence of distinct geneiTi in Australasia
(Australia, Tasmania and New Zealand), in New Britain and in the Malay Peninsula
and Sumatra, rendere it impossible, in the absence of information concerning the
representation of Peripatus on any of the intervening islands, to consti'uct a scheme of
any route of migi-ation in former geological ages, especially as we cannot be certain
whether the centre of distribution lay in the Eastern or the Western Hemisphere,
although M. Bouvier' is of opinion that the American Continent was the original head-
quarters of the Onychophoran stock.
lentil quite recently a peculiar correlation has seemed to subsist between three classes
of phenomena in their relation to Peripatus, namely: (1) the discontinuity of its areas of
distribution : (2) the generic divergence of the different tj'pes ; (3) the method of nutrition
of the young in the intra-uterine development.
It now appeal's certain that this con-elation is neither so simple nor so fixed and
definite as had been supposed. If the method of embryonic nutrition, for example, were
strictly correlated \\'ith the generic divergencies, the selection of any particular method as
bearing a more primitive or ancestral stamp than the rest, might well appear arbitrary-,
since the generic divergence may be taken to depend partly upon the discontinuity of
distribution.
When I firet became acquainted with the blastodermic or trophic vesicle of the
embryos of the New Britain species {Parapeftnpatus novae-britanniae) I was completely at
a loss to offer any provisional explanation of it based upon morphological principles, but
in the course of time I evolved a working h\-pothesis, an account of which was published"-.
Through the kindness of Professor G. B. Howes, F.R.S., I have recently come to the
knowledge of an interesting memoir by Dr H. J. Hansen of Copenhagen, on the stnicture
and development of a primitive insect named Heinimerus talpoides, belonging to the order
Orthoptera'. This insect has been found living in the fur of a West Afiican rat of the
genus Cricetomys : it resembles a wingless cockroach ia supei-ficial appearance but is more
nearlv related to the Forficulina than to the Blattina. as is indicated especially by
the aiTangement of the mouth-parts. The chief peculiarity of Heinimerus lies in its
anomalous method of propagation which isolates it fi-om all kno\\"n insects. It is truly
\i\-iparous (not merely ovoviviparoiis) and in one female Dr Hansen foimd six embryos
at different stages of development, the largest of which, in its convoluted state, measured
2'8 nun. in length and the smallest 1'2 mm.^
Dr Hansen points out that it is thus evident that Heinimerus gives birth to its
voung singly and in succession, not simultaneously; moreover the newly born 3'oung onl}-
differs fi-oni the adult in the less number of joints of the antennae and in lacking
the sexual modification of the posterior abdominal segments.
' Zool. Anz. XXIV. 1901, p. .5'.t.
-■ \Yilley, A., " Tiophoblast and Serosa." A contributiou to the morphology of the embryonic membranes
of insects. Quart. J. Micr. Sc, Vol. 41, 1899, pp. 589—609.
^ Hansen, H. J., " On the strncture and habits of Heiiiimenis talpoides Walk.," Eiitomol. Tidskr. xv.
Heft 1, 1894, pp. 65—94, Pis. 2 and 3.
■• The length of the parent without the cerci was 11-5 mm. [Hansen.]
PERIPATUS. 705
In the four largest embryos which he extracted from the uterus, Dr Hansen
observed a most remarkable feature. From the articular membrane which connects the
head with the protergum, there issued an unpaired nuchal organ, slender at the base
but rapidly increasing in volume, which became lacerated during the removal of the
embryo. " This organ," he considered, " must be in connection with the interior wall of
female genital organs and thus serve the nutrition of the young ones."
While taking out the two smallest of the embryos, which he was able to procuri'
without destroj'ing the specimen, the head was broken off in each case, which seemed
to him to indicate " that also these specimens had been fixed by an organ from the
neck." Remarking upon the probably nutritive function of this nuchal organ, Dr Hansen
does not hesitate to give it as his opinion that the very anomalous mode of propagation
of Hemimerus is of great systematical importance', and marks it as the t}-pe of a distinct
fiimily allied to the Forficulina.
To my mind the nuchal organ of Hemimerus is homologous with the corresponding
trophic vesicle of Paraperipatus novae-hritanniue as described by me^, all that is wanted
to complete the comparison being an account of the actual history of the organ in
Hemimerus in order to demonstrate its relation to the normal embryonic membranes
of the insect.
The other fact which it is my fortune to bring forward in support of the suggestion
that the trophic vesicle here referred to is a formation of primary- or phylogenetic
significance, is due to the recent observations of M. Bouvier of Paris^. In a species of
the South Afi-ican genus Peripatopsis, namely, P. sedgwicki Purcell, M. Bouvier describes
a blastodermic vesicle with nuchal insertion as in Paraperipatus. The vesicle attains
relatively enormous dimensions, measuiing as much as 5 mm. in length in the case of
an embryo of -3 mm. M. Bouvier is of opinion that the species in question is the
most primitive member of the genus Peripatopsis and that the other species of this genus,
<vs regards then- development, are derived from it by atrophy of the vesicle accompanied
by greater simultaneity in the growth of the embryos.
' The expression "great systematical importance" may be interpreted in various ways, but, for my part,
] prefer to take it as signifying great phylogenetic importance.
- Willey, A., "The anatomy and development of Peripatu.^ novae-britaiviiae," Zool. Results, Part i. 1898.
•' Bouvitr, E. L., " Observations sur le developpement des Onychophores," C. R. Acini. Paris, tome 131,
p. 6.52, 1900. JI. Bouvier has again returned to this question (" Sur la reproduction et le developpement du
Pi-ripalopsis blainvillei," op. cit., tome 133, p. 518, 1901) in connection with the Chilian species whielk affords
a transition from the Andicoloas to the South African forms. He finds that the genus Peripatopsis presents
three modes of development: — 1. avec un oeuf tr^s petit et sans annexe embryonnaire (P. bluinvillei) ;
2. avec un ceuf assez volumineux et [avec] vesicule nuquale {P. sedgwicki) ; 8. avec un oeuf assez volumineux
et sans annexe embryonnaire (tous les autres Peripatopsis). In the second category 1 have placed the word
"avec" in square brackets, because, in the original, the word "sans" is inserted by an olivious typo-
graphical error.
That all these deductions cannot be accepted without discussion goes without saying, but fortunately the
issue is fairly clear and an important presentation of the opposite view has quite recently been contributed
by Professor Arthur Dendy (" On the oviparous species of Onychophora," Quart. •/. .Micr. Sc, Vol. Li, 1902,
pp. 363—415, Pis. 19—22). Dendy thinks that the oviparity of certain Australian species which lay large
yolk-laden eggs is primitive, and adduces in support of this contention the consideration that the chorion
which surrounds the intra-uterine embryo in the viviparous species of Peripatiis may be regarded as a vestigial
structure inherited from oviparous ancestors in which a chorionic membrane acted as a protection to the
deposited egg during the early stages of development. It does not follow however that the eggs were primitively
overloaded with yolk, since most so-called alecithal eggs are also protected by a chorionic or vitelline membrane.
706 THE MYSTERY OF THE PEPE.
I desire to take this opportunity of qualifying my former statements concerning
the structure of the endoderm in post-natal and in prae-natal (i.e. immediately prior
to birth) stages of P. novae-britanniae (Part i. Zool. Results, pp. 29 and 30V This
seemed to afford ground for comparison with certain histol}i:.ie phenomena which have
been observed to take place in the intestine of insects and some other animals. In
his recent detailed description of the Malayan genus Eoperipatiis (published in the
Qtuirterly Journal of Microscopical Science (1901), Vols. 44 and 45), Mr Richard Evans
finds no confirmation of the observations which I recorded, and gives reasons for
supposing that the appearances were the result of the method of preservation. After
renewed cijnsideration and re-examination of my preparations I have come to the
conclusion that he is probably right. This qualification has no reference to the
" trophocytes " which I described in an earlier stage {op. cit. p. 22), to which I adhere.
Being desirous, as occasion offered, of acquiring some personal acquaintance with
one or two of the native sorceries I brought my mind, in the Easter of 1895, to
Consent to be initiated into the mystery of the " jjepe." The details had been very
kindly an-anged for me b}" Mrs Parkinson by negotiation with a native petty chief
named Taipuk.
From information which Mrs Parkinson' obtained from the natives, the "pepe" is
somewhat vaguely described as a preparation made by native doctoi-s to be partaken
of by others for the purpose of causing the spirits which dwell in certain trees to
enter theii- bodies for specific purposes. There are several trees which are frequented
by the spirits, notably the banyan tree, " a giau."
The " pepe " is prepared once a year during the south-east season when it is
dry, because in north-west weather the spirits are driven fi-om the trees by the rain.
At the right time, as ascertained by the behaviour of the tides, the man who prepares
the " pepe " scrapes the aromatic bark from certain trees, adds to it Ai-eca-nut and
lime, and «Taps the whole in the " Aier " or betel leaf, making a number of small
bundles which he spreads out in the less frequented parts of the bush. WTien the
" pepe " is laid out in this way, i.e., in the manner of a repast, the people who intend
to partake of it gather round it and the master gives each communicant a small
parcel. They eat their parcel and then fall down in a trance as if intoxicated, and
go to sleep on the spot. When they awake in the morning, they start off by twos
or threes to find the spirit-haunted trees, and to sleep under them until they discover
by the nature of their dreams a tree inhabited b}- a good spirit, i.e., a male spirit,
as the female spirits are reputed bad. The information which they desu'e to obtain
from the good spirit relates to personal ornaments, waistbands made with shells, chaplets,
tattoo-marks, feather-tufts for the head, scented leaves for the neck, dyes for the hau%
pig's bi-istles for the hair, cassowary quills and opossum-teeth for the nose, painting
c>f the body and so forth. Further information is desired upon the important matter
of charms, chiefly of the natiire of love-philtres, fern-seed and rain-making. All this
knowledge is imparted by the good spirits in dreams to him who has taken the "pepe."
The spirits have different names and different functions. One is called " Tutana Vuragit "
' Mrs Parkinson was kind enougli to present me with a description of the " pepe " which she reduced
to writing for the occasion. The general account which I give in the text is largely derived from this document.
THE "pepe" festival. 707
and is described as the everlasting spirit ; another is " Pepe tuna " or the true spirit
of the " pepe " ; and there are others. After a certain lapse of time the " pepe " votaries
gather together in a special " pepe "-house to recount their visions, and then decorate
themselves according to their respective inspirations. Revelations concerning future
events, such as dances, are held in abeyance until the appointed time.
While holding communion with the spirits which dwell in trees, when a man,
under the influence of "pepe," "exempt from public haunt, finds tongues in trees,"
he does not associate with his friends, but keeps in retirement until finally the maker
of the " pepe " gives a great feast for the men, towai'ds the expenses of which each
guest contributes a yard of diwara, and the spell is broken.
I took my '• pepe " in a .secluded spot in the cocoa-nut plantation behind Ralum,
but it was unconvincing, as they were afraid to give me too much " pepe," and I failed
to seek subsequent inspiration under the proper trees. Nevertheless the ceremony of
the " pepe " undoubtedly presents many features which are both quaint and picturesque.
I was expected to contribute to the repast a native dish called " a ku," which
my boys prepared. This consists of the juice squeezed out from the white kernel of
the cocoa-nut, which is boiled in banana leaves by placing heated stones in it. Under
this treatment the milky juice becomes as thick as pomdge, leaving a wateiy residue
which is drained off. The creamy decoction was then tied up in the leaves and carried
to the rendezvous, accompanied by a fish of moderate size, which had also been cooked
and wrapped in leaves. The " pepe " was tastefully arranged on the gi-ound in a circle,
placed upon leaves with the margins artificially coloured red, and, outside these, wreaths
of similar leaves, and also variegated leaves of Dracaena, Croton, and other ornamental
plants cultivated expressly for such occasions by the natives. The whole was protected
by a fane of palm-leaves. After I had paid my initiation fee of two fathoms of diwara
the palm-leaves were removed, the pepe exposed and handed round. When taken, the
leaf with its red-coloured margin is wrapi)ed round the mixture of bark-scrapings and
nuts, and the entire bolus is then swallowed, a taste of ginger predominating. They
had prepared some six or eight leaves, which were distributed among as many persons
including myself, but they said that the whole quantity prepared should have been
eaten by one recipient, in order that its effects might be appreciated. The painted
leaves used at the "pepe" and on other occasions are carefully hidden away afterwards
so that they may not be looked upon by the women.
Shortly after the experience in practical folk-lore which I have described, namel}-
on April 17th, 189.5, Mr Parkinson invited me to accompany him to a place some
miles distant in the bush behind Ralum, where a great dance or " malagSne," called
the '• maravot," was going to be held. It had been convoked by an aged chief oi'
wealth and fame, named To-ragat, the richest and most renowned personage of all the
country round Ralum.
The " maravot '" is an elaborately concerted festival designed for the purpose of
initiating the young men into the tribal usages, and touches the very heart of the
native society of the Gazelle Peninsula. It is not held annually, but at long intervals,
and like all such tribal assemblages of people who do not possess written rccoixls to
give authority to their precedents, its scope will become narrowed and it will sooner
w. VI. 93
708 THE " MARAVOT,"
or later cease to exist. I may therefore esteem myself fortunate in having witnessed
this rare scene under conditions which enhanced its ethnological value and human interest.
It has been fancifull}' suggested by various authors of books of travel that the
Papuans and other races similarly situated are relict types of the Stone Age, since in
their ignorance of iron and metals generally, they are dependent upon stones and
organic productions (shell, bone, wood, fibre, leaves) for theii- martial and industrial
implements. It is therefore impossible to avoid a feeling of regret when we reflect
upon the sure approach of the relentless wave of destruction which will engulf all
these neolithic rites.
The accessory incidents of the " maravot," the mise-eii-scene and appurtenances, if
described in too great detail would appear sordid, perhaps even repulsive. But if it
be regarded sympathetically as a manifestation of racial sentiment, its importance
becomes at once evident, and one who has recently received his baptism of '' pepe "
should be able to approach the "maravot" in the proper spirit'.
At the place of meeting, on the occasion of our visit, there was a lar'ge oblong
enclosure fenced in by tall, opaque screens of plaited palm-leaves so as to exclude the
gaze of the women to whom it is " tambu." Nevertheless, at a little distance from
the enclosure there was a " monstrous regiment of women " bringing large bundles of
food, called " magit," wrapped up in banana leaves. Beyond a few armlets and necklaces
of C«sc«s-teeth (which are worn alike by both sexes) the women had not decked
themselves out, and they presented a rather monotonous sea of faces, as they sat on
the ground smoking, chatting and chemng, while their male relatives attended to the
more serious duty of dancing in the sanctuary.
The men (with the exception of the great chief To-ragat, who came quite plain)
had decorated their persons with all the ornaments which the country yields, theii"
toilette being completed by a head-ch-ess consisting of a tuft of white tail-feathers of
the domestic fowl. Many of them had painted the body, face and hair with red, white
and black native pigments, and some were actually green with smeared lichen. The
wealthier individuals wore precious collars of Ciiscus-teeth, nose-pieces and chaplets.
At one end of the enclosure there were some remarkable mural paintings repre-
senting figures of reptiles with human heads, forming an appropriate mounting for the
piece ; at the opposite end, faced by the natives while dancing, there was an exhibition
of coils of diwara belonging to various chiefs, this in itself being a sufficient proof of
the solemnity and sincerity of the occasion.
When advancing to take their places in serried ranks within the sacred precincts,
where they presented the appearance, viewed from an elevation, of a bed of white
feathers, the men formed processions in single file, chanting as they walked and carr^dng
coloured leaves and carved devices. One very large figure representing a departed chief,
' In the dictionary of the New Britain dialect by the Bev. E. H. Rickards, to which I have already
made reference, the "maravot" is defined as the ceremony and dancing initiatory to the " igiat " (pronounced
"ingiat"), which is, as far as I coidd gather, a secret fiaternitv which most of the men join iu order to
settle their accounts in this world and the next to the best advantage. I think it might be possible to
compile a connected account of the native economy of this part of New Britain based upon the " igiat,"
elucidating all the ceremonies and periods wliich precede membership and all the privileges and powers
which ensue.
B^^'-M-* =^^ • '**■ >■ "» Tar- ]& ' 1 J
r* "v 4 ■^mV■>« i i'fyi Ml Wk- 1 ir. n
i *■: 1 '»'
^l>■ -wRi ^i ^#-Y ^'^1.. ^^4^ - ^ ^
Fig. 8. Screen-paintings within the maravot enclosure (New Britain).
Fig. g. Natives who took part in the maravot. The central figure is a petty chief |Taipuk> who hke
his two neighbours wore a collar of Cuscus-teeth.
7V» /nrr pttge 708.
THE " MARAVOT. 709
had a liead consisting of a human skull covered with dark plastic material (" a tita ")
and was also adorned with a C'«scKS-necklace. As the procession passed along, the
women raised a howl, perhaps bewailing departed spirits. The life-size human emblem
and another smaller one were placed in fi-ont of the diwara, facing both the |ierformei-s
and the altar-piece at the far end.
Some of the dancers brought very curious carvings representing a tongue proti-uding
through an open mouth. These were placed upon a rough log bench immediately in
front of the altar-piece, and I inferred that they symbolised spirit-tongues, but have
no information on this point.
Thf actual " nialagene," in the densely crowded enclosure, consisted of an in-
terminable medley of unintelligible but well-rehearsed incantations delivered by one or
other of the assistants in a high-pitched voice and followed by a chorus of responses,
shouts and chants, all in unison, accompanied by dancing and some comedy. Each
distinct dance was inaugurated by the entii'e company, at one moment, blowing out
fi-om their mouths a cloud of betel juice into the air, an operation known as " puruk.''
As I have already indicated, the bulk of natives, constituting what I may term the
chorus, faced the diwara \vith their backs to the altar-piece, but the leaders, first
among whijm was the venerable To-ragat, faced the crowd and appeared to direct the
performance.
Some of the young novitiates, sons of chiefs and valiant men, gifted wdth superb
figures and well-chiselled features, commanded instant admiration ; but the older con-
gregants improved on acquaintance.
Among the lowei- orders of marine animals to whicii I de\'oted some attention
\\hile dwelling on the shores of Blanche Bay, two or three species of Ascidians deserve
special mention on account of some striking circumstance of habit or structure.
I have described one of these under the name Styeloides eviscerans\ It has a \ery
characteristic form, colour and encrusting method of growth which distinguish it ft-om
all the remaining somewhat numerous species which I observed during my entire
expedition. It also differs from other Tuniciita by the readiness with which it throws
out its entrails, in a manner analogous to the evisceration of some Holothurians, shortly
after removal fi-om its natural surroundings. The extrusion is eff'ected by successive
contractions of the atrial siphon, and if the tunicary be cut open there is found to
be no trace of lesion inside. Moreover the animal appears to be perfectly healthy and
normal externally after having got rid of its viscera, and both the liuccal and tile
atrial siphons remain sensitive to mechanical and chemical stimuli.
I found a few examples of this species attached to the lower surface of stones oft'
the south-west shore of Vulcan Island at a depth of one-half to one fathom. The
general colour is a dull reddish-lirown, more pronounced near the apertures, the margins
of which are dark red, interrupted by four light streaks indicating the characteristic
quadripartite nature of the siphons. Individuals occur singly and associated in groups,
in which case they may be so intimately united by fusion of the test-substance that
one would be tempted to assume that the group represents a colony of ascidiozoids
' Willey, A., " Letter.s from New tiuinea on Xautilut and some other organisms." Qituit. J. Micr. Set.,
Vol. .Hll, IH'.IT. p. Uil.
93—2
710
STYELOIDES EVISCERANS.
budded from a parent stock. There is however, so far as I could ascertain by dissection,
no organic union.
Why one species should act in the way described while another attached perhaps
to the same stone does not, is a question not easily answered, but the fact affords
a .special case of self-mutilation or autotomy, a phenomenon of wide distribution in the
animal kingdom, varying greatly in its manifestations.
Fig. 10. Group of seven individuals of Styeloides eviscerans, represented as lying attached to the surface
of a fragment of stone. In the large individual to the right the digestive tract is indicated in process of
extrusion through the atrial aperture, a. Anus with frilled margin, i. Intestine, br. s. Branchial sac.
e. Endostyle. h. Foreign organisms attached to test. (From Quart. J. Micr. Set., Vol. 39, 1896.)
The genus Styeloides was established in 188.5 by Dr Sluiter' for a species found
b}' him near Batavia, which appeared to be normally destitute of a branchial sac.
Subsequently he returned to the subject and suggested that the branchial sac anfl
intestine are lost in the adult jjeriod only'^
' Natuurk. Tijdsclir. Nederl. Imh, Bd. 4.5, 188.5, Styeloides abranchiata. Sluiter.
- Sluiter, C. Ph., "Tunicaten." Semon's Zool. Forschiingsreisen, v. 1895, Styela solvens, p. 182.
RHOUOSOMA HUXLEYI.
711
More recently Dr Sluiter has accepted my observations as affording an explanation
of the absence of the branchial sac in Styeloides, and he thinks that this generic name
should lapse in favour of Polycarpu'.
Another noteworthy Ascidian which I found in Blanche Bay on a reef-patch off
Baravon, is called Rhudusoma hiuieiji, and was described under the name Pera hixlei/i
in 1862 by Dr J. D. Macdonald, who obtained his material from the Bellona ReeP.
The peculiarity of Rhodosoina is this, that the upper ])ortion of the test fits like
a lid upon the rest of the tunicary and can be raised and depressed like the valve
of a lamellibranchiate Mollusc. Compared with those bivalve Molluscs where one
valve is fixed to the substratum, the action of the lid of Rhodosoma which protects
the siphons bears a striking analogy to that of the freely hinged valve of the former.
It is, in fact, a good example of parallel evolution, a phenomenon of wide occurrence
and profound significance, but not always easy to recognise. By what may be termed
a singular coincidence, the late distinguished Professor H. de Lacaze-Duthiers", who was
one of the first to publish a full account of Rhodosoma, was at great pains to demon-
strate the supposed molluscoid affinities of the Ascidians, and the discovery of a species
of this peculiar valvate genus confirmed him in what mcist naturalists will now admit
to have been an interesting and instructive error.
*^s*;iM^->
Fig. 11. Rhodosoma hu.vleiji ; front view with operculum raised; side-view with Ud closed.
(Semi-diagratumatic, partly after Macdonald.)
Finally there is a third Ascidian which is abundant in Blanche Bay and
neighbouring localities (New (iuinea, etc.) and claims notice on account of its size and
I Sluiter, C. Ph., " Tunicaten von Sud-.\frika." Zool. Jahrb. Si/sl. xi., 1898, p. 5. It is not known
whether the viscera can be regenerated, but Dr Sluiter thinks he has found indications of a regeneration of
the endostyle in Styeloides ahranchiata.
- Macdonald, J. D., "On a new genus of Tunicata (Pera) from the Bellona reefs.'' •/. IJiin. Hoc. Zouh,
VI. p. 78, 1862. I am not acquainted with any later record of this species.
* Lacaze-Duthiers, H. de, " Sur un genre nouveau d'.\scidien {Chevreulhis)." Ann. Sci. Nat. {')) iv. ISO."),
p. 293. The name Rhodosoma, which takes priority, was introduced by Ehrenberg in 1828.
712 SPAWN OF CEPHALOPODA.
brilliant orange hue but more especially on account of the remarkable fragmentation of
the dorsal tubercle (orifice of the subneural gland into the branchial sac) which appears
as a large number of in-egularly placed apertures occupjong a triangular area. This very
curious species is the Ascidia aurata of Quoy and Gaimard, who gave a coloured figure
of it in their work on the zoology of the Astrolabe'.
My enthusiasm for the Ascidians resulted in temporary disablement for the best
part of a week, owing to the combined action of the sea-water and the heat of the
sun, which so blistered my back that I experienced a complete ecdysis of the dorsal
integument.
On May 6th, 1895, I took over b\- purchase a stout cutter-rigged sailing-boat built
by Ah Tam, the enteqjrising Chinaman of Matupi, with whom I had been negotiating
for some time. I was delighted \\'ith the new sails and the bamboo-boom and with the
prospect of being able to trawl at will'-.
There was, however, ground for considerable anxiety as there was more or less
constant danger of the boat being stolen by deserting labour-recruits. One such theft
occurred about that time with disastrous consequences, involving the death fi'om exposure
of the owner who went in pursuit.
Among my various devices designed to secure the eggs and young of Nautilus
I employed a tangle-di'ag, a bar of iron with long hempen tangles hanging fi-om it.
which was sunk in deep water in places frequented by Nautibis and left for a vai-jing
number of days with a buoy at the surface. I obtained the spawn of other Gephalopods
by this means, but not that of Nautilus.
Another obvious method of dealing with this intractable creature was to bait and
sink fish-baskets in the usual way, and to leave them do-ivn for several days instead
of raising them at the customary time. This method was based upon the principle that,
while it began to seem hopeless to think of acquiiing the eggs in any other way than
by inducing Xautilus to breed in confinement, it was not necessary to look at the captive
specimens at once. In this way I sought to avoid the shock of bringing them to the
surface before they could have a chance to settle down to the work of o\'iposition.
Pre\'ious experience iiad led me to anticipate that the pro^'isioning of the cages after the
exhaustion of the bait would be accomplished automatically by the entrance of Crustacea
and soft-shelled Echinids. Nautilus duly entered the traps, but in spite of sanguine
hopes the experiment did not succeed, owing probably in part at least to various
hindrances, such as the fragile construction of the cages, the inroads of conger-eels and
the buffetings of other large fishes in the depths.
A further hostile element which has to' be reckoned with when operating in deep
water is the periodical ^asitation of gales of ivind or lialf-gales, especially during the
north-west season. These will often mercilessly dispel all one's hopes and contrivances.
1 Quoy et Gaimanl, " Vo.yage de I'Astrolabe," Zoo/. T. iii. 1834, p. 604. .\tla<< (Mollusques, etc.) PI. ^1.
fig. 3. Dr Sluiter (Tunieaten iu Semon's Forschuni/sreiseii, BJ. v., 189-5) has described this species under the
came Stijela {Poli/rariKt) piwiinionodes and was the first to note the pecuUar condition of the dorsal tubercle
(cf. Sluiter, op. cit., Atlas Taf. x. figs. 1—3).
- I procured more material with a small beam-trawl, which I took out with me from Plraiouth at the
recommendation of Mr J. T. Cunningham, than with the dredges, among my prizes being two very fine
Polychaet worms belonging to the genera Psammolyce and Pahmjra.
" MALIRA. 7 1 3
" Paraparaii " and " malira " are local ordinances i-elating to marriage. The former,
as I understand it, corresponds to a betrothal, but does not involve the appearance of
the prospective bride. If a man is in a position to maiTy, he makes or observes
" paraparau," going into retirement for a time, organising festivities and leaving the
question of a bride to the last. It is defined by Mr Rickards, the author of the
dictionary from which I have previously quoted, as the native custom of concealment
during the preparation of marriage.
"Malira" is a term applied to substances employed as love-charms, and it also
appears to be the more general name for a marriage festival or fair'. At all events
towards the end of May (1895) a "malira" was held at Karavia close to my house.
All through the night on the eve of the fair, natives were coming in from the bush,
shouting at intervals in a loud monotone, blowing conchs and beating garamats. When
day came the company sang chants.
At a "malira" which I attended at a place called Pal-a-kulau (literally, house of
cocoa-nuts) behind Karavia, the natives sang chants in a specially erected spirit-hut
which was surmounted by feathers, rattles and coloured leaves and filled with carvings
(if human figures and birds. Scores of natives attended, marching in procession and
canying wooden effigies of pigeons suspended from long switches. One man came
wearing a skull-mask ("a lor") made from the fiicial portion of a human skull plastered
over with " tita," hair and beard being ingeniously glued on, in true semblance of
humanity. The carvings were all on the flat except the pigeons. There was also
a distribution of " bue " (betel-nuts) and diwara, but the latter was presented to certain
people only. The food, contributed from different quarters, was wrapped up in banana
leaves and was all paid foi- with diwara. The dancing, in which the women joined,
consisted in circling round a " garamat " and was accompanied by singing.
In illustration of the working of " malira " in its quality of love-philtre I may quote
the following story which came to my knowledge. It seems that a man and woman
who are "nimuna=" with one another cannot eat or even carry food in each other's
presence, this abstention being a mark of respect. When a man desires to pay court
to a woman he will lie in wait for her as she is walking through the bush and will
then throw a " malira^ " at her. If she knows who the man is through previous
understanding and if she likes him, she will pick up the charm and put it in her
basket. But if she intends to reject her suitor she will roundly abuse him and may
lodge a complaint with her husband, who Avill promptly demand " tambu " fioui the
offender. This is obviously a fruitful source of blackmail, and a case of the kind
hiippened near Maulapao-* shortly before I left New Britain, a man being accused by
a woman who was his " nimuna." The woman stated that as she was walking through
1 The women are soil-tillers as well as wives, and I think the "malira" is a kind of fair analogous to
the old sessional fairs in the provincial towns of England where farm-labourers were hired.
• - The term " nimuna " signifies the relationship between a man and his wife's parents and with those
who are "tamana" with her (Bickards). ••Tamana" is the relationship between father and offspring and
between the latter and the father's brother.
3 Consisting of a piece of bamboo or leaf with son\ething wrapped up in it, perhaps some bark-scrapings
from a particular tree with a little lime mixed in.
■• The name of the site upon which Mr Parkinson's house stands.
714 NATIVE FANCIES.
a patch of the long grass called kune or kunai, which is used for thatching houses, she
heard something rustle in the gi'ass, but in searching for a " raalira " was unable to find
one. Whereupon ^Nith true native logic she made a long detour and finally found her
victim hiding away with a cemple of cocoa-nuts, as he knew his " nimuna " was passing
through the neighbourhood and he could not let her see him \vith food in his hands.
In consequence of the woman's assertions her husband demanded ten fathoms of diwara
fi'om the alleged offender although there was no proof whatever against him. If the
woman heard anything at all in the grass it was most likely a lizard and she concocted
her story on the strength of it. The last I heard of the affair was that the parties
had submitted the matter to Mrs Parkinson's arbitration.
In the course of my relations with my assistants I became acquainted with several
of the minor traits of the native character.
Sometimes they fall ill and ascribe their malady to the influence of " puta." They
are very careful to conceal or destroy all waste-products of their food, an admirable
custom, but if they fail to do so by inadvertence, an enemy can collect the refuse and
work mischief with it by sorcery. This branch of sorcery is called " puta " and one of
my men, To-warabur, fell a \'ictim to it, according to his own account.
When the}' see a log of wood or a bamboo-stem, floating vertically in the water
and bobbing np and down, as often happens, they are fiightened and filled with
veneration for it, calling it " a tuk." On one occasion we passed such a drifting
bamboo rod upon the tojj of which there were some accidental marks so disposed as
to convey a grotesque resemblance to the human face. One of my boys, To-iraran, fled
behind the mast in terror of this ill-omened taberan or demon. I then leamt that
a fonner assistant of mine, a " koloata " or man of the bush, called To-bokat, who had
fallen sick and returned to his village, had traced his affliction to the retribution due
to him for having innocenth' hurled some stones at one of these vagrant logs.
Once I thought I had purchased the fidelity of two bearded men of Davaun but
they disappeared during the night. I went to Davaun the next day to demand an
explanation and was told that To-palavur's mother was sick and had said " verj' good
they no go belong whity man." This affords an illustration of the working of the native
idea of the " kubak," according to which the fi-iends and relatives of a sick person who
have slept within the same house or enclosure as the j^atient cannot spend a night
elsewhere without causing a turn for the woree.
In the middle of July I left Karavia to take up my quarters in a recently vacated
trader's house at Baravon on the other side of the bay, where there is good anchorage
during the south-east season. I stayed here for some time, the chief objection to the
place being that it was overrun by rats, which manifested the utmost impudence and
indifference to the j)resence of a white man in their midst.
In August I paid a visit to the Credner Islands, called locally the Pigeon Islands,
of which Nanuk is the smaller and Palakuvur the larger. These uninhabited islands are
purely coral-formations, covered %\'ith vegetation, surrounded by reefs with a passage
between the islands and a beach of white calcareous sand. On a subsequent occasion
I procured here numbers of the interesting Enteropneust, Ptychodera ruficollis, but this
time I accidentally incapacitated myself for serious work by biting into an uncooked
TALILI BAY. 715
taro. I had anchored on the reef at Naniik for the night, it being high water, and
intended to sleep on the boat while my boys lighted a fire and made themselves
comfortable on the beach, cooking taro for themselves and for me. Dviring the night
it rained and thundered, and thinking to console myself with a taro I gi-oped in the
dark amongst the litter of ropes and gear, picked up one of these excellent fruits of
the earth and made a lusty bite into it. It proved to be uncooked but I recovered
from the dire effects \vithin the twenty-four hours.
In the middle nf August a note was brought to me from Mr Reuben Buckley,
a trader quartered in the Malagunan district quite at the other end of Blanche Bav,
saying that Nautilus was also found off the north coast of the Gazelle Peninsula.
Accordingly the next day I paddled over to Malagunan and was conducted overland
to a place called Matakabang on the north coast, the road between these two villages
being flanked on the east by the heights of the North Daughter. From Matakabang
we pulled across to a district at the head of Talili Bay called Unakamkambi, where
there was another trading station tenanted by Mr MacNicoll. Here I made arrangements
for traps to be set for Nautilus and next day returned to Blanche Bay to raise up
some coral blocks which I had previously sunk in a selected spot in about 80 fathoms.
On examining the coral trees I found niunbers of the large Ostracod crustacean which
has been described by Mr Stebbing under the name Asterope arthuri, but no signs
of Nautilus.
The next morning I set sail for Unakamkambi, rounding Praed Point and Cape
Stephens. I spent the best part of a month at this place, having found the conditions
for jjursuing my search for the young of Nautilus almost as favourable in the south-east
season in Talili Bay as they were in Blanche Bay, but not more so ; in the north-west
season I think it would be impossible to operate in Talili Bay. Near the eastern limit
of Talili Bay there was a Catholic Mission station, administered by Pfere Bley, and at the
western point a Wesleyan Mission station, presided over by Rev. J. A. Crump, at the
■ village of Kabakada adjoining Unakamkambi.
I have made occasional reference to certain New Britain usages according as I had
opportunity to observe or to take part in them, and it is impossible to leave this island
without a word concerning two of its most characteristic and well-known institutions, the
Dukduk and the Tubuan. They are cognate organisations ordained for the glorification
of departed chiefs and to promote the circulation of diwara, although this brief statement
does not exhaust the sum of their functions. It appears that the Tubuan is supposed to
die periodically and to be born again. I once witnessed a most interesting representation
of the resurrection of the Tubuan, which took place opposite to my house at Barawon
near the village ruled over by an important chief named To-kinkin.
At midnight, in the month of July (189-5) a couple of Tubuans fi-om Matupi, escorted
by a fleet of forty or fifty canoes, arrived at To-kinkin's place. They had brought with
them a carved human emblem representing a personage whom they called To-ririg,
doubtless a great man of former days. This circumstance suggested to me the possibility
of the dukduk and its subsidiary tubuan being a means by which the history of the
w. VI. 94
716 TUBUAN AND DUKDUK.
tribe is hcanded dowTi fi-om one generation to another^ Throughout the rest of the night
the natives uttered awe-inspiring howls at frequent intervals. After suniise they took
to their canoes again, the two Tubuans in separate canoes, and the entire flotilla pulled
out to the middle of the bay, where they came to rest. The Tubuans thereupon
executed an elegant dance with due restraint in their respective canoes, apparently
illustrating theii- I'egeneration fi-om the sea in the manner of Aphrodite-. The setting
for the performance provided by the natural features of the bay was most picturesque
and impressive, and the effect was heightened by the chant which was intoned by the
assistants.
Tubuan Chant.
JS^Wi
' tx i ' ^ \ ^^ ^ — 6^—1 — • ti»- i ' — ix — i ' ^^-^
' ^ ^ \ — — ti*— p^
The more important Dukduk is dressed like the Tubuan in the bird-costume, the
chief difference being that the conical head-dress of the former is much longer than that
worn by the Tubuan. The Dukduk observance may be a very elaborate ceremony^, but
for my part I saw some dukduks on the beach at Matupi on one occasion only. I was
informed that they had been engaged by an old chief who expected to die but recovered.
On several other occasions while I was in New Britain a Tubuan was to be seen
dancing along the beach skirting Blanche Bay, shouting to scare the women. Sometimes
the Tubuans are engaged instead of the Dukduks, I think on account of the smaller
expense, to celebrate the funeral rites of a deceased chief. None were present at the
only native funeral which I attended, but another funeral was taking place elsewhere
at the same time, at which two Tubuans were employed.
If the seven principal features of New Britain life which I have briefly touched
upon be enumerated in the following order, pepe, maravot, igiat, paraparau, malira,
tubuan, dukduk, we shall gain I think a fairly accurate insight into the sequence of
ceremonial observances in this wonderful island.
Towards the end of Sef)tember (189.5) the " Ivanhoe," a topsail schooner of
152 tons, Capt. James Steele, unexpectedly arrived at Herbertshohe from British New
Guinea on a trading cruise. She was to return to Cooktown by way of New Guinea
and after long consideration I determined to continue my quest in the neighbourhood of
Milne Bay, knowing well the importance of keeping on the move while engaged in such
work as that to which I was committed. I took leave of my kind fi-iends at Ralum with
' Compare Graf von Pfeil, " The Duk Duk and other customs as forms of expression of the intellectual
life of the Melanesians." Rey. Brit. Ass., 1896 (Liverpool), p. 939.
- The costume of the Tubuan, consistiug of a tall conical mask topped by a plume of white feathers
and passing below into a heavy stack of leaves reaching from the shoulders to the knees, is intended to
represent the figure of a bird, and what is known of bird-symbolism may suffice to justify the above com-
parison, although I have no actual information on the subject.
'■> Cf. Meyer, A. B., and Parkinson, B., Album der Papua-Tijpeii, 1894. Pis. 13 and 16.
MILNE BAY AND HYGEIA BAY. 717
/^
many expressions of regret and sailed away on board the " Ivanhoe " on October 12th.
With a fair wind we should have reached Saniarai or Dinner Island, the headquarters
of the Government in the Eastern Division of British New Guinea, within four days,
but contrary winds, squalls and calms delayed our arrival until October 2.5th. Here
I lost no time in procuring a small decked cutter, the " Mizpah," which at once
provided me with shelter and means of conveyance, although I was not so much in
need of the latter, having brought my boat from New Britain. But the need of
sleeping accommodation decided me to acquire the " Mizpah " and, on the whole, I had
no reason to regi'et the purchase.
The shells of Nautilus are found in great numbers among the islands which compose
the eastern archipelago of New Guinea, and it was this information which induced me
to ascertain whether there might be any prospect of meeting with a more suitable
locality in these waters. It is a curious reflection that although Nautilus ranges over
a large part of the Indo-Pacific Ocean fi-om the Philippines to the Fijis, at no point
within the sphere of British influence' is it available for practical investigation. Stra}'
specimens may be caught here and there but it is only to the Philippines, Moluccas,
Bismarck Archipelago or Loyalty Islands that we may look for the final solution of
the mystery surrounding the propagation of Nautilus.
The natives of the country round Milne Bay and of the neighbouring islands are
not great fishermen and do not make elaborate fish-baskets, so that they contrast very
noticeably in this matter with the men of New Britain. Perhaps the reason for this
may be that animal food is more plentiful in the bush in New Guinea than in New
Britain, where pork may not be eaten by members of the tribal fraternity of the igiat.
Whatever the explanation may be, the fact remains that while the natives may be
seen fishing with a seine on the reefs, they neither employ the larger traps nor do
they know how to make them.
I commenced operations in the D'Entrecasteaux Group, putting up a temporary
house on an uninhabited coral islet called Karuana, close to the important island of
Sanaroa-. This was a delightful spot and a capital site for a biological station, as
the surrounding seas contain numerous coral-patches.
I exjjlored Hygeia Bay on the coast of Fergusson Island. On this island the
natives were rather a wild-looking lot, such of them as came down to the beach.
They can-ied cosmetics and charms in bamboo^ tubes, and some of them wore head-
ornaments or chaplets consisting of a fringe of yellow feathers (fi-om the crest of the
cockatoo) rising from a border of red parrot feathers, closely resembling a similar
ornament used by some Indians of the Amazons region.
I put down a basket here but the resulting captures of a small viviparous shark,
Crustacea, etc. offered little encouragement for Nautilus. There is an extensive reef at
' With the possible exception of the Fiji Islands from whence, so far as I know, no preserved specimens
have been sent either to Australia or to Europe, in spite of the existence of circumstantial accounts of the
methods employed by the natives to catch them.
- Welle Island on the Charts.
' Bamboos do not occur in all parts of New Guinea and its dependencies, and those which grow on
Fergusson Island are a feeble variety in comparison with the bamboos of New Britain and elsewhere.
94—2
718
AMPHISII.E STRIGATA.
the north-west corner of the bay and good anchorage on a sandy bottom at the south-
west comer where there is also a deep freshwater creek not marked on my charts.
I took three soundings across the mouth of Hygeia Bay in a northerly direction
showing 59 and 66 fathoms of water, but at the third sounding the line was bitten
across, probably by a hungry shark, and the lead was lost.
Having injured my foot on the coral at Karuana my principal boatman, Suopa,
made me a poultice by scraping some raw yam with a shell and cooking it in
a frying-pan; he told me to put a little sugar in it and apply it to the foot. I did
so and I think it undoubtedly assisted the inflammation and reduced the pain.
Frequently my boys would break out into song and sometimes with theii- clear
natural voices the effect produced was quite charming. Once when I was busy in my
cabin my skipper started a spontaneous anthem in honour of the south-east wind
(" bomatu ") and was soon joined by the little cook in unison.
Hymn to the south-east wind.
Boma-tu - na Bo - ma - tima
Si - wai sru - ba
Bagi - gi-no Bo - ma-
tuna
i
A
T=^
\J
m.
wai gu - ba - raia
-P-Bz3^
§
=1-=);
Bagi - gi-no Bo - ma - tuna.
W;:q3p-r^:
;s^==^=
Among the reef-patches at Karuana I made the acquaintance (if a remarkable
species of fish, the Amphisile strigata^ of Dr Giinther. This species lives in small
shoals of about half-a-dozen individuals and swims about with rapidity in a vertical
position, cleaving the water with its razor-shaped bod}^ I was greatly astonished when
I saw these spectral fishes flitting about and I pursued them with much perseverance
in my dinghy, but although I came up %vith them several times they doubled back
with such deftness that they completely baffled my attempts to catch them with a
pole-net. Subsequently I obtained specimens at Lifu in which I recognised my Karuana
friends. The tiny mouth is placed at the summit of the pointed snout and is only
capable of imbibing microscopic nourishment.
At the eastern extremity of the mainland of New Guinea there is the famous
inlet of Milne Bay, at the sight of which I was tempted to believe that all things
were possible in its untried depths, but so far as Nautilus is concerned I met with
no better fortune here, after trapping and trawling for a fortnight, than in Hygeia
' Kindly identified by Mr G. A. Boulenger, F.R.S., from specimens taken at Lifu.
NATIVE DEVOTION.
719
^-
^i'
Bay. It is true that a single specimen of Nautilus umbilicatu.^ was picked iij) froui
the surface of the neighbouring sea and reached my hands hiter in a mangled condition.
From this circumstance it would seem possible
that systematic search on an extensive scale
with unlimited equipment would discover a
Nautilus ground in New Guinea waters. But
from all I can hear, it is likely that the
Philippine Islands will prove to be the vantage
ground for this investigation'.
An old pearler whom I met in Milne Bay
on board his cutter informed me that he did
not think Nautilus lives in Milne Bay, but the
shell drifts into the bay from the windward
from such places as Teste Island, Brooker
Island and the Du Chateau Islands", although
some natives said they had taken it in the bay
long ago.
Shortly after leaving Milne Bay I was
Sony to lose the services of mj' native skipper,
Suopa, who was a cheery fellow as well as a
handy man. Knowing that it was my ultimate
intention to leave New Guinea in order to seek
out some more likely place, he one day offered
to follow me wherever the four winds might
take me, not caring even if it should so fall
out that he never returned home. I was rather
touched by this assurance of devotion, the more
so because I knew that it would be contrary
to regulations to accept his offer of service
away from New Guinea, but I was quite unprepared for the reaction which set in the
very next day when he demanded money and freedom, and, not content with these,
began raking up old scores as a reason for lea%'ing.
Next day I sailed east, in the direction of the Conflict Group, with a crew of
three mop-headed youths, including a little fellow whom we called " Cookie," but whose
real name was Taliwanedi, a native of Sanaroa.
The distances between the numerous groups of islands into which th(> eastern
division of Papua is subdivided, are not very gi'eat, but the opposing winds, currents
and calms render navigation uncertain and sometimes precarious even to a seasoned
mariner.
\\\
Fir,. 12. Shoal of Aiupliisile stiifiatti in attitude of
swimmiD". They are representeil as if swimming
from right to left. (Drawn by Mr Edwiu Wilson.)
On the 14th of January, while nearly becalmed and tfrifting within sight of the
low-lying western islets of the Conflict Group, a shell of Sepia, often called the cuttle-
' Cf. Dean, Bashford, "Notes on Living Santilus." Amer. Xalural. xxxv. pp. 819 — 837, lilOl.
- I had intended to visit these islands, but the imminence of the south-west monsoon in this region
prevented me from getting farther east than the Deboyne Group, except at a risk of being wind-bound.
720
CTENOPLANA.
bone, came floating by, borne along by a t\vo-or-three-knot current tn the north. One
of niy boys who knew my interest in flotsam, jumped into the dinghy with jjraiseworth}-
dispatch and secured the shell.
It had been afloat long enough to allow many kinds of larvae (Crustacean,
Anthozoan, Annelid) to attach themselves to it whether as a 'temporary resting-place
or as a permanent foundation. I placed it in an enamelled basin with sea-water,
whereupon many of the organisms detached themselves and swam about in the dish.
My attention was soon attracted by some small flattened disc-shaped bodies about
one-thii-d of an inch in diameter which could both creep and swim. As they crawled
along they jerked out a pair of milk-white tentacles having a worm-like motion of
their own, and quickly retracted them, repeating the process continually. My joy was
complete when I recognized in these small creatures the very remarkable genus
Ctenoplana of which only one specimen had ever been seen before, namely in 1886
by Prof KorotnefF, who found it in the plankton or drift-fauna off the coast of Sumatra.
Although cramped for space in the little cabin of my cutter I had the necessary
appliances for making a cui-sory examination of anything which required immediate
attention and was thus able to make some additions and emendations to the account
furnished by the original discoverer, and I considered that my re-discovery of Ctenoplana
made amends for a great deal of trouble and disappointment.
Fig. 13. Ctenoplana ; (1) from above in attitude of creeping with tentacles extruded ; (2) and (3) side-
views in attitude of swimming. The first two are the red form, the third is the green variety which contained
male gonads; t.o. po.sition of orifice of the sheaths of the pinnate tentacles. (After Willey, Quart. ■/. Micr. Sc.)
Ctenoplana is one of those puzzling creatures which rack the brain of the contro-
versial morphologist in his futile effort to arrive at a provisional settlement of a point
CTENOPLANA. 721
in phylogeny. When it is creeping ulong the bottom of a dish or, back downwards,
along the surface fihn of water, it looks for all the world like a Polyclade Turbellarian.
The fitful extrusion of the opaque white tentacles which have sei-pentine intrinsic
movements of their own, reminds one involuntarily of the behaviour of the Nemertine
with its proboscis. On the other hand, when Ctenoplana undertakes to swim, it does
not do so by the graceful skirt-like evolutions of the Polj-cladf, but it assumes a helmet
or Pilidium shape, depressing the two halves of the body until they assume a vertical
position like the flaps of a bonnet, and then a set of organs, called ctenophoral plates,
which are utterly foreign to Turbellarians but distinctive of a group of hyaline pelagic
organisms, the Ctenophora, come into play.
The ctenophoral plates which, as I have said, are the swimming organs of Ctenoplana,
occur as eight small oval areas on the dorsal surface. Each plate is beset with a
limited number of transverse rows of powerful cilia agglutinated together at their bases
so as to form a membrane upon which the cilia are inserted like the teeth into the
back of a comb.
These characters seem to indicate that Ctenoplana occupies a central position midway
between the pelagic Ctenophora and the littoral Plathelminthes. This, however, is the
simple view of the matter and there are many practical obstacles to its adoption.
Some zoologists contend that Ctenoplana is to be interpreted as a specially adapted
creeping Ctenophore, but as a result of my observations, which unfortunately left many
questions still unanswered, I advocated its ordinal autonomy, chiefly on the ground of
its possession of distinct male genital ducts opening to the exterior on the dorsal
surface of the body^
The principal features of interest in the organisation of Ctenoplana, features which
cannot fail to mark it out for special discussion whenever the interrelations of the
lower orders happen to be adjudicated, may be summarised as follows: —
I. Abstract positive features.
a. Dorsiventral differentiation.
b. Biradial symmetry.
c. Main axis passing between the median ventral mouth and tiie median dorsal
sensory apparatus.
II. Abstract negative features (involved in the preceding but worthy nf special
tabulation).
((. Absence of antero-posterior differentiation.
b. Absence of ambidextral differentiation (there is no light and left).
III. Concrete positive features.
a. Ctenophoral plates.
b. Semi-opaque, pigmented integument.
c. Paired pinnate tentacles (Greiftentakeln).
d. Paired aboral, ciliated, sensory papillae.
' Willey, A., "On Ctenoplana." Quart. J. Mici: Sc, Veil, xxxix., 1897, pp. 323—342. IM. 21.
722 CTENOPLANA.
e. Ciliated ectoderm on ventral surface.
f. Male gonads with tiuiica propria.
g. Male genital ducts opening to the exterior on the dorsal surface.
The fact of biradial symmetry means, of course, that there are two jjlanes of
symmetry which divide the body respectively into two equal halves. One of these
planes passes through the muscular pinnate tentacles, and is called the tentacular plane
of Gtenoplana and of the Ctenophora generally. The other plane cuts the tentacular
plane at right angles, and coiTesjjonds to the so-called stomachal or gastral plane of
the Ctenophora. CoiTesponding with these two planes we have the two axes, namely,
the tentacular axis and the gastral aads.
Arnied with a few new fects, one is strangely tempted to enter the lists of
liiological speculation and take one's chance in the meUe, or in other words, to make
some addition to the discussion concerning the relation of the above-named planes and
axes of bu'adially symmetrical animals to the planes and axes of bilaterally symmetrical
animals such as Turbellaiians and Annelids. The chief question which fixes the attention
of the observer is this : — Is it the tentacular plane or the gastral plane which is
homologous with the sagittal jjlane of a bilateral worm ? or, to put it more brusquely.
Is it the tentacular axis or the gastral axis which corresponds with the oro-anal axis
of a Nemertine or an Annelid ?
It might be and I believe it has been contended that there is no gi-ound for
comparison between the axes of a Ctenophore and those of a worm, in which case
the question jDropounded above has no point. . Freely admitting that this may be true,
there is still room to suppose that it may not be true. Assuming therefore that one
or other of the biradial planes must represent the sagittal plane of higher forms, I
give it as my own conviction that the tentacular jjlane of Gtenoplana about which the
aboral ciliated sensory papillae are disposed in paLred gi-oups as shown in Fig. 13,
coincides with the sagittal plane of a bilateral Turbellarian, Nemertine, or Amielid.
Much as the prominent pinnate tentacles of Gtenoplana^ simulate transversely-paired
organs, they are not more paired than are the ciliated sensory tentacles of the same
animal. Both these organic systems, however, are paii-ed about different planes and it
is to this circumstance that the discussion which I have outlined is due. I simply
desire to emphasize what I believe to be theoretically sound, namely, that if the
above-named planes are held to be respectively comparable, then the tentacular plane
of Gtenoplana and of the Ctenophores- represents the sagittal plane and the pinnate
tentacles occur in this plane, like the proboscis of the Nemertines. Either there is
no correspondence at all or the tentacular and sagittal planes are homologous, but to
my mind there can be no question of the gastral axis of Gtenoplana corresponding
with the antero-posterior or oro-anal axis of the bilateral forms.
Whatever the truth may be, Gtenoplana remains a remarkably interesting tj'pe and
no zoologist could encounter specimens of it without experiencing a momentary thrill
' The same applies to the Ctenophora in a less obvious sense. For discussion of the axes of Ctenophora
the reader should consult Prof. Chun's Monograph, Die Ctenophoren des Golfes von Neapel, 1880.
- This is the view, as concerning the Ctenophora, which was first held by Professor Chun, but he seems
to have made a subsequent retractation.
CTENOPLANA. 723
of satisfaction. Of the four specimens which I obtained three were mottled with green,
the fourth with red pigment; the former were rather larger than the red variety and
differed from it somewhat in shape. The red form also differed in minor details from
the original tj-pe Gtenoplana kowalevskii Korotneff. Nevertheless, with added experience
of specific divergencies, I am now inclined to think that my Ctenoplana rosacea is
co-specific with Korotneff 's type, and it seems quite likely that the green form is
nothing more than a variety or phase of the same species.
All the green individuals contained male gonads with their genital ducts, but I
was unfortunately unable to satisfy myself concerning the female gonads. It should be
noted that great care and the application of very special methods are requisite for the
preservation of Ctenoplana. It is well known that some of the Ctenophora dissolve
away before one's eyes in certain fluids, and although Ctenoplana does not vanish in
this way, the contraction of the tentacular musculature is apt, in healthy specimens,
to cause a lamentable shrinkage.
In his recent account of the Ctenophora' Mr G. C. Bourne places Ctenoplana and
the allied genus Coeloplana in a special order of Ctenophora Tentaculata, which he
names Platyctenea, at the same time declining " to attach very much importance to
the resemblance between Ctenoplana and the Polyclada." Mr Bourne further comes to
the conclusion " that the Turbellaria, the Nemertines and the Ctenophora are descended
from a common ancestor which is most nearly represented by the larva of Stylochus"
[a Polyclade worm]. This is mainly in accordance with the views which I expressed
in my paper, and Mr Bourne quotes the Pilidium larva of Nemertines in his
comparison, as I also did. Moreover, the term Platyctenea is undoubtedly preferable
to the term which I suggested, Archiplanoidea, as an ordinal name, but I am unwilling
to refuse any importance to the latter name. I intend it to embody the proposition
originally laid down in its main principles by Prof. Chun, that the bilateral Plathelminthes
generally have originated from a biradially symmetrical ancestor, and further to express
the possibility of the diphyletic origin of triploblastic animals.
After much labour against the tide we managed to get to a temporary anchorage
off the westerly islet of the Conflict Group, intending to make the lagoon before dark,
but failing to find the passage we spent a squally night tacking about. Towards the
evening a remarkable procession of transparent pelagic animals, Ctenophora of several
different kinds, including the ribbon-shaped Gestus, drifted past the islet. These creatures,
as I have said, are most difficult to preserve, as they have the peculiar property of
undergoing instantaneous disintegration and dissolution .under certain conditions of
treatment.
Next morning I had quite lost my bearings and suddenly found myself heading
straight for the reef, but I got the jib and sail down just in time to avoid a collision.
It was not until the following sundown that we found the passage and anchored in
the lagoon. Just about this time the south-west wind began to blow and I found
the lagoon with its loose sandy bottom a most imcomfortable anchorage. In this region
' A Treatise on Zonlogy, edited by E. Kay Lankester, I'art ii. chap. vii. 190i). The Ctenophora by
G. C. Bourne, M.A.
W. VI. 95
724 DEBOYXE GROUP.
the two predominant winds are the south-east and the south-west monsoons, the latter
corresponding with the inclement season which in fact begins to set in towards the
end of January. The numerous islets surrounding the Conflict Lagoon, which is upwards
of 20 miles in length, are uninhabited, the natives merely using this as a half-way
house on their long canoe voyages. At the time of my \-isit, as I have said, the
south-west squalls commenced and, increasing in severity, my two anchors persisted in
dragging, so that on the fifth day there was no other course open to us than to get
clear of the lagoon which had become a veritable trap, if we wished to avoid the
ignominious piling up of our craft on the reef. After being foiled in several attempts
we emerged successfully through a narrow passage nearly opposite to the one by which
we had entered the lagoon. I think it must have been blowing quite half-a-gale and
the sea was running high, but the ilizpah's thin boards held together and we spent
the night tacking and drifting in the usual manner. On the following day although
the wind had moderated I had again lost my bearings and was at my wits' end to
know which course to set in this reef-bound sea, when we espied another cutter two
or three miles ahead and decided to follow her lead. In due course our unconscious
pilot disappeared through a passage in a distant reef and I headed straight for this
point to the best of my belief, but on arrival was confronted by a continuous reef
which seemed to stretch on either hand as far as the eye could reach from the mast-
head. By following up the reef, though labouring under total misapprehension as to
the identity of the various islands around us, we suddenly found ourselves in a -svide
passage which, as we subsequently ascertained, led us into the magnificent lagoon of
the Deboyne Group, at the northern end of which stands the high crescentic island
of Panaieti, overshadowed by the loftier crest of Misima to the north-east.
Inside the Debo}Tie Lagoon there is another hilly island called Panapompom, oS
which we anchored and set about purchasing food, our stores having run out, but only
bread-fruit was procurable. The natives of Panaieti are in the habit of crossing over
to the south-east side of Panapompom in order to cultivate gardens of yams, sweet-
potatoes, pumpkins and bananas, which however do not form a conspicuous feature in
the scenery as they do in the Engineer Group and other more westerly islands.
While at anchor in the lagoon I put do^vn a small dredge, taking it out in the
dinghy for some distance, then sinking it and hauling in ^vith the hand-^\•inch on
board the cutter. I was rewarded for my pains by finding in the sand with which
the dredge became filled a species of lancelet (Amjihioxus) belonging to the genus
Asymnietron, which had only been discovered a few years previously by Prof. E. A.
Andrews' at Bimini in the Bahamas.
The family of the Branchiostomidae, which includes all the known species of the
lancelet, occupies such a peculiarly central position in the system of animal life, and
its distribution is so local, that it is always satisfactory to ascertain a new fact
concerning either its organisation or its range-. I was not a little surprised to find
' Andrews, E. A., "An TJndescribed Acraniate : AiymmetTon lucayanum." Stud. Johns Hopkiits Univ.,
v., pp. 213—247, 1893.
= Willey, A., "Zoological Observations in the South Pacific." Quart. J. Micr. Sc, Vol. 39, p. 219, 1897.
Amphioxus belcheri has its southern limit in Moreton Bay, Queensland, and its northern in Japan. It has been
LANCELET,
725
a representative of this West Indian species in the Louisiade Archipelago and subse-
quently in Sandal Bay, Lifu, although it is a fact that there is a strong Caribbean
element in the fauna of the Indo-Pacific. Still it is interesting in consideration of
the fact that two other species of the same family belonging to different genera,
Amphioxus belckeri and Epigonichthys cultellus, occur in Torres Strait and the Moluccan
Sea but have not been found in the West Indies.
Fig. 14. Asymmetroii ciiiiddliim (probably a sub-species of A. lucayamim). A number of the oral cirri are
united together by an intertentacular membrane ; the uotochord is continued as a urostyle behind the muscle-
segments. Magnified about 7 times.
At the time of my visit, as I was informed by the resident Wesleyan missionary,
the Rev. Ambrose Fletcher, the population of Panaieti numbered 466 men, women, and
children, who live in a series of villages on the south side of the island facing the
lagoon. The island is rocky and the soil unproductive, so that food is often scarce.
The natives, however, are great sailors and canoe-builders and cover long distances by
means of their huge mat-sails. More than half the population may be absent from
the island during the south-west season, in search of food, pigs, yams and so forth.
The houses are well built and have a characteristic shape, somewhat resembling a huge
inverted whale-boat, often with a verandah-like platform in front. In every village
there was a small bar of wood supported upon a couple of uprights and bearing
numerous lower jaws of pigs to show what the natives had eaten, the skeletons
of former feasts which could be contemplated in times of stress. The uprights were
often surmounted by a cocoa-nut, reminding me that in New Britain the husks of
cocoa-nuts were wont to be displayed in a similar manner. It woidd appear that these
people exist largely upon the " bare imagination of a feast," and at this time of the
year (January) food was very scarce, bread-fruit and nuts called "siaia" being the chief
commodities, while hunger can always be staved off by chewing the areca nuts which
they obtain from Misima. The " siaia " nuts are roasted in the fire or cooked in
clay-pots and have an agreeable taste. Another species of nuts called " dausia " has
to be soaked for five days in sea-water either before or after being cooked, in order
to remove its noxious properties and render it edible.
The piles upon which the houses of the Panaietians are raised, are fitted with
large circular wooden discs perforated in the centre to admit the post ; they are called
" panapana " and are intended to prevent the incursions of rats into the dwellings.
Tattooing is not practised at Panaieti, but the teeth of the women are blackened
and the ears pierced upon arriving at puberty. Should a young girl join the mission,
the old women of her village will urge her to leave the white man (" dimdini "),
recorded from Singapore by the late Mr Bedford. The lancelet of the Bass Strait (Iletcroplcuron hassanum),
as I ascertained from specimens in the Australian Museum, Sydney, kindly .shown to me by Mr Thomas
Whitelegge, ranges north as far as Port Jackson.
95—2
726 INDUSTRIES OF TUBETUBE.
saying, " Your teeth are not blackened and your ears are not pierced, no man will
marry you."
After having been kindly entertained by Mr Fletcher, who insisted upon my
acceptance of a supply of cqffee and tea, I started to leave Panaieti shortly after
sunrise on January 30th, with a light wind. Towards noon a northerly breeze carried
us through the passage between the Torlesse Islands and the Panaieti reef. There was
a lively tidal rip in the passage, and just as we emerged from it the wind ceased
and the tide turned simultaneously, as so often happens, and we drifted back into the
lagoon, where in the pale moonlight we soon found ourselves scraping the surface of
a submerged reef I did not know where the next tack would bring us, but it happened
that a welcome land-breeze came to our assistance, enabling us to make the passage
once more during the night, and a couple of days later we reached the island of
Tubetube in the Engineer Group.
The population of the Engineer Group was distributed as follows, Tubetube 350,
Kuriva 260, Naruaruari 120. The natives are famous sailors and tillers, but poor
fishermen. When mourning for relatives the women blacken their bodies, wear long
leaf petticoats reaching nearly to the ground, a mass of cords round the neck and
a necklace called "kokoana" of the large shells of Ovulum ovum ("dunari"). Only the
women are tattooed all over the chest and abdomen and often upon the face as well.
The large canoes of Tubetube are made at the Woodlark Islands (Murua) and
are decorated with bird scrolls at the ends and fish-representations on the hull. Wooden
bowls in use here are also made at Murua, but the wooden scoops for baling out the
large canoes are manufactured at Tubetube as well as at Murua. The clay bowls of
Tubetube constitute a staple article of trade with other islands, more especially with
Dobu ; they are extensively moulded by women on the three islands of the Engineer
Group, being shaped by hand without the aid of pottery whorls, by superposing circular
bands of clay ; of course they remain unglazed.
After another flying visit to Dobu and Kaniana, where I had built a pile-dwelling
on a former occasion, I returned to Samarai, from whence I made a voyage (accompanied
by Mr C. H. Walker, with whom I stayed for a few days at Matadona on the western
side of the China Straits) to the island of Suau, which forms the south cape of New
Guinea. In several villages along the south-east coast I noted isolated instances of
the round discs fitted on to the pillars of the houses as at Panaieti. I was told
that there is a story current of the Savaiians (inhabitants of Savaia, a large village
on the mainland not for from Suau, situated at the head of a bay, across the mouth
of which stretches a formidable barrier reef) having fitted out an expedition to fight
the Panaietians some years previously.
By this time I had become convinced that it would be useless to spend more
time in searching the coasts and islands of New Guinea for Nautilus, in spite of the
frequency with which the shells are cast upon the reefs. I therefore decided to go
still farther east, either to New Caledonia, Fiji, or the New Hebrides, in any case by
way of Sydney.
I spent my last somewhat fever-stricken fortnight in New Guinea under the
hospitable roof of the Rev. C. W. Abel, of the London Missionary Society, on the small
i. rf" .^v S ,
,/1^^>^- I
Fig. 15. Shore-scene at Panaieti showing pile-dwellings with rat-guards on
the posts ; a canoe is hauled up on the beach.
Fig. 16. A potter at work on the island of Tubetube ; she is wearing a necklace
of the shells of Ovulum ovum.
To face page 72t5.
PTYCHODERA FLAVA. 727
island of Kwato opposite to Rogeia, within easy distance of Samarai, and left on board
the S.S. Titus on March 28th, bound for Sydney by way of Sudest and the Solomon
Islands, the full enjoyment of this interesting voyage being marred by intermittent
attacks of fever.
We passed Sydney Heads and entered Port Jackson on April 20, 1896. I spent
upwards of two months at Sydney, where I was accorded working space in the Biological
Laboratory of the University by Professor \V. A. Haswell, F.R.S., and was thus able to
examine some of the material which I had collected, every ftxcility and kindness being
showered upon me by Professor Haswell and by my friend Dr J. P. Hill. I was also
made free of the excellent library of the Linnaean Society of New South Wales, by
the courtesy of Mr J. J. Fletcher, M.A., and received valuable information from the
Director of the Australian Museum, Mr R. Etheridge Jr. and from Mr Charles Hedley
and Mr Thomas Whitelegge. I learnt from Mr Hedley that Nautilus macromphalus was
definitely known as the " Isle of Pines Nautilus " and this practically decided me to
wend my way to New Caledonia. I therefore left Sydney on July 2, on board the
Messageries Maritimes S.S. Tanais bound for Noumea, making the passage through the
marvellous barrier reef of New Caledonia on the fourth day out from Sydney. As some
days must elapse before I could secure a passage to the Isle of Pines I made arrange-
ments through the instrumentality of gentlemen to whom I had brought introductions
from Sydney to accompany the Breton pilot, M. Le Ravallec, in the pilot-cutter to the
lighthouse on the He Amedee, about twelve miles out from Noumea, where the pilots
await the arrival of steamers.
On the shore of Amedee I obtained my first specimens of a remarkable species of
Enteropneusta, Ftychodera Jlava, which I afterwards found more plentifully at the Isle
of Pines and also on the weatherside of Lifu, that is, on the side opposite to that on
which Sandal Bay is situated. From the point of view of my own experience the
observation of this species in the living condition, subsequently enhanced by the study
of its organisation, threw a flood of light upon the structure of this group of burrowing
" worms," which respire by means of gill-clefts, and it seemed to me that Pt. jlava
should be regarded as the tyj^e of the Enteropneusta from a morphological standjKiint
as it actually is upon chronological grounds, an opinion to which subsequent examination
lent additional support.
It is a remarkable feature of morphological inquiry that one worker should find so
many points of comparison between distantly related groups where another, often a very
eminent authority, sees nothing of the kind; others again engaged upon similar m-
vestigations but starting with different premises ignore utterly the existence of groups
of animals which seem to bear upon the matter. For example, I am ready to affirm
with considerable confidence that the unpaired hollowed "roots" which pass between the
medullary canal of the collar region and the epidermis, where they join the intra-
opidermal fibrillar plexus in Ftychodera flava and in allied species, belong to the same
category, in other words, are homologous with the pineal outgrowths of the primary fore-
brain of Vertebrata. This is a conclusion which may possibly be supported by fiirther
observations, but can hardly be submitted to the test of experiment, and must therefore
ever fail to convince the rightly sceptical.
728
PEOCHORDATA.
The diagrams here reproduced are founded primarily upon conditions prevailing in
the larva of Amphioxus, which are of the utmost complication as soon as one approaches
Fig. 17. Diagrams to elucidate some suggested comparisons between an Enteropneust (A), an Ascidian
larva (B), and a Craniate embryo (C).
A. p. proboscis ; p.p.c. proboscis pore-canal opening close to the anterior neuropore which leads into
c.v. the collar nerve-tube; e.p. epiphysial roots; s. stomochord; m. mouth; c. collar-region; g. gut; t. trunk-
region; d.n,t. dorsal nerve-tract; a. anus; py. pygochord.
B. f.o. organ of fixation; e. endostyle; m. mouth; a.n.c. anterior neurenteric canal which, together
with s.n.g., subneural gland, constitutes the neuro-hypophysis ; c.v. cerebral vesicle; sp. medullary tube
communicating with the gut by means of p.n.c. posterior neurenteric canal ; n. notochord.
C. ep. pineal body or epiphysis cerebri ; f.b. m.b. and h.b. fore-, mid-, and hind-brain ; hy. hypophysis
cerebri ; th. thyroid gland ; other letters as above.
them in a morphological spirit. In the Prochordata (including for the present occasion
the Enteropneusta, Tunicata, and Cephalochorda) the three principal canal systems of the
body, namely, eoelomic, neural, and alimentar}', open in various ways to the exterior in the
anterior region, where they form an intricate association of apertures which I have formally
designated as the anterior trematic complex.
The follo\\ing table will possibly be useful to future investigators who may ventm-e
to ■i\Testle with this problem.
PROCHORDATA.
729
Tabulation of structures concerned in the formation of the anterior trematic complex of the
Prochordata, consisting of organs which are probably homogenetic and of their connections which are
presumably related through metaleptic succession or substitution.
I. Enteropseosta.
II. Tdnicata.
III. Cephalochobda.
Proboscis
with a head-cavity or proboscis
coelom communicating by a coe-
lomic orifice with the proboscis
pore-canal (or end-vesicle) which
opens externally by the proboscis-
pore in the vicinity of the anterior
neuropore.
Organ of fixation
with a caviti/ in which mesen-
chjTiie-cells circulate ; probably a
degenerate eoelomic cavity but
neither blastocoel nor haemocoel ;
it is a deuterocoel sui generis.
Rostrum
with a head-cavity which becomes
divided in the embryo into right
and left portions, the former con-
stituting the rostral cavity and the
latter the praeoral pit of the larva
and the ciliary complex at the roof
and sides of the oral hood in the
adult. The praeoral pit repre-
sents, inter alia, the eoelomic
orifice of I., being removed from
dorsal surface by intervention of
notochord ; physiologically it is
comparable with the dorsal tubercle
of II.
Proboscis pore-canal
which, as indicated above, opens
externally near to or in common
with the anterior -iieuropore.
Neuro-hypophysial canal
or duct of the subneural gland.
It arises in jirimary continuity
with the wall ijf the cerebral vesicle
with which it remains in com-
munication in the larva by an
intracerebral orifice ; at the other
(oral) end it opens into the vesti-
bule of the branchial Siic by an
intrahuccal orifice which gives rise
to the dorsal tubercle of the adult.
Olfactory pit
into which the anterior neuropore
opens in the larva.
Ventral mouth
of larva and adult.
Dorsal mouth
of larva (with which the nem'o-
hyjiophysial canal communicates)
becoming the buccal siphon of
adult.
Sinistral mouth
of larva (separated together with
the praeoral pit from association
with the neuropore by intervention
of notochord) becoming the median
ventral velum in the adult, the
innervation of which, according to
the discovery of Professor Van
Wijhe, remains sinistral.
Having obtained the necessary permission from the authorities of the French
administration I left Noumea on July 18 on board a local steamer for the Isle of Pines,
accomplishing the journey of 65 miles in nine hours. The southern coast of New
Caledonia is studded locally with forests of Araucaria, and the Isle of Pines is a hilly
satellite of the larger island, and likewise affords some exquisite coast scenery. It is used
as a penitentiary establishment, the native population being confined to a reservation on
one side of the island, the most picturesque side, where it is controlled by the Mission
authorities under the direction of Pere Lambert, who is a renowned conchologist. I paid
730 ISLE OF PINES AND MARE.
several \'isits to the native who was the nominal chief of the tribe, but he failed to
understand my ■ivishes and showed me no hospitality. I made an abortive attempt to
reach his heart by means of a present, which he accepted, and sent a man in brass
buttons to me on the following day to know what it was for.
Altogether I found the Isle of Pines an unsatisfactory place in which to pursue my
special and peculiar avocation, and the eighteen days which I spent here were only
relieved by two circumstances, namely, the opportunity of studying Ptychodera flava
which I found at low tide in sandy rock pools on the coralline platform at the base of
the cliffs where the breakers expend their fury, and secondly the companionship of my
native servant, an important chief from Lifu who had been exiled from his native place
for drunkenness accompanied by violence. A man of forty with a distinguished face and
bearing, speaking fair English and better French, he served me well in return for his
food and one franc a day. He would have made an ideal " roi de Lifu " but for his
addiction to the \dle vitriol which does duty for liquor. He answered, temporarily, to the
name of Eugene, and in the course of conversation he strongly urged me to make for
Lifu rather than Mar^, although I had been thinking of the latter. Fortunately his
counsels prevailed and I left the Isle of Pines on August 5 in a cutter manned by Mare
islanders. We anchored for the night at a point called Gadji, at the northern end of the
Isle of Pines. Here there are extensive reefs, mud-flats, quicksands and sand which is
strongly elastic, rebounding to the tread like india-rubber, imparting an iinusual and
somewhat exhilarating sensation. There are also numerous islets clothed with vegetation
and undermined in a characteristic manner by the action of the sea\
The next morning at 4 a.m. we left Gadji with the rising of the moon and had
a fair wind to Mare, where we arrived at sundo^vn. There is some difficulty in landing
here as the rock-bound coast drops sheer into the sea, and the rudeness of the rocks
simply anticipates the astonishing inhospitality of the natives. It is true I presented
a dilapidated appearance, especially after walking over the scraggy limestone surface of
the island, and had it not been for the good offices of a petty chief called Wainani, who
had been a fellow passenger on the cutter, I should have been in a still more desolate
condition, being already mistaken for a libere. The Government representative on Mare
was the Commandeur, M. Journot, who entertained me at luncheon on my way to the
great chief Neisselin at Netchi. With the best will my friend Wainani none the less led
me into a trap, by which I became exposed to the merciless and humiliating diatribes
of Neisselin's wife, delivered in excellent English. The population of Mare consists of
Catholics and Protestants, and the latter have for some years been left without a white
shepherd, but they conduct their o^vn services with a gusto which must be heard to be
appreciated.
Besides the usual beehive huts they have white plaster houses which are rather
oppressive. I boarded with a native called Ito Pupu and after prayers went to bed.
Next day he informed me that his father had partaken of white man. I had agreed
with a tall handsome bearded native called Waiyowara, the owTier of a cutter, for my
passage to Lifu, the arrangement being that he would call for me at a certain place
and time. He was not punctual, and such was my anxiety to leave this island where
' Similar mushroom-shaped islets occur in the Fijis and have been described and figured by Mr J. Stanley
Gardiner in the Proc. Camb. Phil. Soc, Vol. ix. Part 8, 1898; see p. 459.
SANDAL BAY, LIFU. 731
I had suffered so much mortification, that I shipped in a small half-decked cutter called
the "Marie Therese," with a dangerous native called Pedro as skipper. The boat
manifested a pronounced tendency to capsize, and had actually turned turtle on a previous
occasion when several Isle of Pines men were drowned. We did not clear the point of
Mar^ without a mishap, as the boat, failing to respond when we were coming about,
was driven upon a reef-patch, where she toppled over on her beam-ends. By poling and
shoving we managed to escape on the crest of a wave and put back in a leaky condition
to Netchi.
My only course now was to reopen negotiations with Waiyowara, and by dint of
allowing him a just measure of reproach and other concessions he consented to land
me at Chepenehe on Lifu, where I had been informed a trader, Mr James Wright, had
a station. After reporting myself upon arrival, to the French Resident, I amxnged to
stay in a small outhouse belonging to Mr Wright and take pot-luck with him for my
food. I stayed here for some eight months, in the course of which some Nautili which
I had succeeded in keeping alive in captivity deposited their eggs in my cages. Most
unfortunately these eggs proved to be infertile, an eventuality which the marine zoologist
rarely expects to encounter, though it would not surprise an ornithologist.
The wanderings which I have described in the foregoing pages were all directed to
one end. I wished to find a locality where Nautilus could be both captured and kept
prisoner in a moderate depth of water, in four or five fathoms instead of forty or fifty.
Here on the shore of Sandal Bay, Lifu, I had come upon the ideal gi-ound for which
I had been searching.
The rocky shore at this part of the bay is riddled with a labyrinth of creeks,
crannies and blowholes. In calm weather some of these natural tanks present a tempting
appearance as if constructed for the benefit of a vagrant naturalist. I soon found
however that the wash of the tide, exerting a powerful suction through the narrow
holes and crevices, rendered these pools quite useless for my purjDoses. Nevertheless the
thought occurred to me that it would not be a matter of insuperable difficulty, given
proper equipment and organisation, to construct a marine reservoir with concrete walls in
which refi'actory animals might be successfully reared, by being accorded a larger measure
of liberty than is possible even in the best fish-trap. Such a tank could also be made
to serve useful puqjoses, as for example the breeding of Holothurians from which trepang
or beche-de-mer is jirepared. Unless the eggs of Nautilus are destined to be taken
in a fertile condition at Negros in the Philippines I can offer no other suggestion than
the above for the benefit of future naturalist- venturers.
Almost any kind of animal bait may be used to attract Nautilus to the fish-traps.
Accompanied by my man Saono, an excellent woodman, boatman and diver, I made
frequent excursions into the bush to procure land-crabs, which live in holes in the ground
beneath stones and logs. To the experienced eye their presence can be known without
turning over a stone, by the neatly cleaned and levelled threshold made liy the crabs
at the entrance to their retreats. When the stone is removed the crab may be at the
top, in which case it scampers away, or it may be in the hole, which is perhaps a foot
long but only a few inches below the surface. The females of these crabs psiss into
the " berried " stage in November and December when they troop down to the sea to
w. VI. 96
732 EGG-LAYING OF NAUTILUS.
deposit their larvae in the water. Their migration takes place about the time of the
fructification of the ferns, when also the yams which have been planted for the ensuing
season begin to shoot up. At this time the}', as well as Birgus, are taken on the
shore by torchlight in hundi'eds, and are cooked and eaten.
Another bait which may be used when others feil consists of crushed sea-urchms,
especially the Heterocentrotus (Acruckidia) mammillatus, which abounds on the reef-patches
and is also eaten by the natives. The best bait is made by cooking the large pra^\^ls or
langoustes, Palinurus (also Scyllarus), which especially frequent the weather side of the
island. When ready they are pounded to a pulp and then wrapped up in the dead
fibrous spathes of the cocoa-nut tree, and placed in the basket, to which they impart
an irresistible bouquet.
Having observed that the newly captured Nautili adhered with great force by means
of their tentacles to the sides of the vessels in which the}' were placed, I fixed some
boxes in the cages. They attached themselves to the boxes but made no other use of
them, and one day a huge conger eel effected an entrance into the trap and brought
serious dissension into the household. To avoid a recurrence of such a disaster we
closed up the entrance with sacking. On the next morning there was a curious white
object, looking at first sight like some part of the axial skeleton of a bony fish, adhering
to the sacking. Ui^on remo^'ing it we found that it had been tightly fixed to the sacking,
and very soon I realised that it was the first egg of Nautilus which rewarded my gaze.
After two years of anxious groping in the dark it may be imagined what a thrill passed
through my marrow, destined to be quenched when I found during the course of the
following weeks that all the eggs which I attempted to rear were infertile.
I kept my culture cages at a depth of about three fathoms and fed the company of
Nautili three times a week, going out on Saono's raft for the purpose.
The Lifuan rafts are well-constructed and seaworthy (see Fig. 5, p. 699). They are
worked by sculling with a long, flattened pole, which passes through a hole in the centre-
board at the fi-ont of the raft, and are sufficiently wide and buoyant to bear the weight
of two men and a large fish-trap between them. Provided that equilibrium is maintained,
a rather difficult matter in a chopjiy sea, the worst accident to be feared is the snapping
of the pole. The position of the fish-traps, which are sunk to depths var}'ing from three
to eighteen fathoms, is ascertained by dead reckoning, buoys not being employed. They
are dimly seen at sixteen to eighteen fathoms on a calm day, and when it is desired
to raise them they are skilfully secured by means of a wooden hook, which is lowered
fi-oni the raft and passed through the meshes. In the " thingit " (the native name for
the fish-traps) in which I kept a stock of Nautilus I had a door made, through which
I was able to get bodily into the cage on board the raft and manipulate at leisure.
I had several of these incubators and tried all methods, pairing off the Nautili in some,
associating them in companies in others, manufacturing dark recesses in sackcloth, all
to no purpose. The eggs were all infertile and often simply consisted of the empty
capsules without any vitellus inside. Generally, however, the eggs contained a healthy
sheiTy-coloured vitellus, which either suffered no change when left for a fortnight or
commenced to addle. It was on December 5, 1896, nearly four months after my arrival
in Lifu, tliat I obtained my first egg and, according to promise, presented Saono with
-V
Fig. i8. Two eggs of Nautilus macromphalus from above, x 2.
{Photographed at the I'jiii'frs'ity of Sydney by Mr Uraiit.]
To /ace pdijc 732.
ASTROSCLERA. 733
twenty-five francs. Near the end of January, 1897, we had a visitation of wind, probably
the trail of a hurricane, which played havoc with my baskets, and cost the lives of
some sixteen Nautili. Shortly aftenvards I lost another " thingit " containing six pairs of
Nautili in forty fathoms, and on March 8 left Lifu in a small schooner, bound for
Noumea, calling at Uvea on the way.
Apart from Nautilus I obtained many valuable accessions to my general collection
at Lifu. A large proportion of Sandal Bay is flanked by terraced limestone cliffs from
seventy to a hundred feet in height. The base is excavated by the sea, as in the case
of the mushroom-shaped islets which I have previously mentioned, and halfway up the
face of the cliff there is an exactly similar excavation. From the base of the cliff the
water deepens rather rapidly to about forty fathoms, and then there is a sudden drop,
showing the existence of a remarkable submarine ledge of dead, shaly rock. I dragged
along this forty fathom ledge with a grapnel in the hope of recovering one of my
" thingits," which had disappeared, and it was here that I found some peculiar calcareous
growths which I preserved, although I had no inkling of what their true nature might
be, suspecting them rather to be Foraminifera. In Mr J. J. Lister's hands they have
proved to represent a very interesting form of calcareous sponge, Astrosclera, offering
a curious superficial resemblance to the mesozoic Pharetrones'.
I also described a curious flatworm^ from Lifu under the name of Heteroplana,
but the generic properties which I ascribed to it are open to another interjiretation.
The shape of the body was peculiar, symmetrical in front, not quite symmetrical behind,
unlike an\i;hing else I had seen.
Examination with lens and microscope in the li\'ing state revealed the fact that
the organs of the left side, princij)ally the intestinal diverticula, were suppressed. I found
the worm in a hole on the lower side of a coral block^ and on account of its large
size, serai-opaque orange colour, characteristic shape and other peculiarities, gave it the
above name. It has been suggested that Heteroplana is a phase in the regeneration
of a mutilated Polyclade worm. If this be so I cannot account for its very definite
external form, but at any rate the genus must remain in abeyance until more light is
thro\\Ti upon this subject''.
The population of Lifu, like that of the other lioyalty Islands, is now entirely
Christian, being divided between the Catholic and Protestant communions. The two great
hereditary chiefs of the island, Ukenesoe and Bula, who formerly waged incessant warfare
with each other, are now the respective champions of the rival creeds. The social
organisation is somewhat communistic so far as this is compatible with the existence
of chiefs, and it sometimes happens that even such valuable propert}- as a horse is
readily jaelded up to the first importunate acquaintance who casts envious eyes upon it.
But of course a return present will be exacted sooner or later. If I gave something
of value, tackle, food or money, to my man, it was quite a chance if I did not find it
shortly in the possession of somebody else in whom I had no particular interest.
' See Appendix.
- I collected a number of Polyclade worms which I have sent to Prof. A. Lang at Zurich. For notes
on oviposition of Polyclades cf. Willey, A., "Letters from New Guinea," Quart. J. Micr. Sc, Vol. xxxix.
'■' Repeated and laborious search failed to eUcit any more specimens.
* Cf. C. Rina Monti, "La rigenerazioue nelle Planarie marine." Mem. Ut. Lombardo, xix. 1900.
96—2
734 SECOND VISIT TO NEW BRITAIN.
Unless some epidemic should come to decimate the population, the future welfare
of the Lifu natives seems to be insured by the poverty of the land. No primary rocks
occur at any part of the island, which consists entirely of coralline limestone with patches
of arable soil, capable of producing a certain amount of food to support a population of
about 7,000, which number appears to remain fairly constant.
I finally left Noumea for Sydney by way of Esafate or Sandwich Island in the New
Hebrides on March 28th, 1897. A few days after my amval for the second time in
Sydney, I despatched the following cable to Professor Alfred Newton, F.R.S., Chairman
of the Balfour Managers : — " No stages, shall I spend rest of year New Britain ? " to which
I received an affirmative reply. Accordingly I set about making preparations for a fresh
campaign in New Britain, the principal addition to my apparatus being a nest of seven
large hencoops, which were made according to my instructions at the Blind Asylum in
Sydney.
I now pass on to the concluding episode of my voyage. I landed in New Britain
for the second time on June 16th, 1897, and on the following day one of my former crew,
To-lara by name, came to see me and agreed to serve mth me again. This gave me an
important start; I revisited my old haunts in Blanche Bay without delay, and having
fitted out my Sydney cages, I stocked them with Nautilus-coivpies as soon as I could
procure them. Very shortly I commenced to find the eggs of Nautilus pompilius
attached to the sacking round the sides of the cages, hardly distinguishable fi-om those
of iV. macromphalits at Lifu. My hopes revived, but I soon found that the eggs fared
no better in Blanche Bay at a depth of 30 — 50 fathoms. No development took place,
and in course of time the vitellus commenced to undergo liquefaction and decomposition.
I continued my efforts, at the same time accumulating collateral material, until the
following September, when I bade adieu to New Britain, and thus terminated my long
quest.
735
APPENDIX.
Mr J. J. Lister, F.R.8., has favoured me with the following additional note, "On the
mineralogical character of the skeleton of Anirosclera," which he has received from Mr Hutchinson
of Pembroke College, Cambridge.
"A short time after the publication of the description of the mineralogical characters
of the skeleton of Astrosdera wUleyana, an important paper was communicated to the
Mineralogical Society by Miss Agnes Kelly, entitled "Conchite, a new form of Calcium
Carbonate" {Min. Mag., Vol. xii. p. 363, 1900). In this paper the authoress claims to
have established the existence of a form of calcium carbonate distinct from calcite and from
aragonite, and to have shown that all calcium carbonate of organic origin, formerly believed
to be aragonite, really belongs to the new variety.
"The specific gravity of conchite is given as 2-830 — 286.5. It is uniaxial and negative.
Its birefringence is less tiian that of calcite, and the maximum value obtained for its index
of refraction 0)0= 1-661 is less than either p or y of aragonite.
"Miss Kelly's conclusions have however been questioned by Brauns {Centralhl. fiir Mill.,
1901, 134) and by Vater (Zeit. fur Kryst., .xxxv. 166), who maintain that conchite is essentially
the same as aragonite.
"Meigen (Centralhl. fiir Min., 1901, 577) has recently shown that aragonite and calcite
in fine powder behave diflerently when boiled with cobalt nitrate solution, the former turning
reddish-lilac, while the latter remains white or becomes yellowish. G. Panebianco {Rivista
Min. Ilal., xxviii. .5 — 12) confirms Meigen's observations as regards aragonite but finds
that calcite turns sky-blue. Astrosdera when submitted to this test gives the colour
characteristic of aragonite. The conclusion drawn from the specific gravity determination,
that the skeleton of Astrosdera is not calcite, is thus satisfactorily confirmed. Its properties
as given in my previous description agree with those of conchite, but until the independence
of conchite and aragonite is more definitely established, the mineral element present iti
Astrosdera may best be regarded as aragonite.
A. HUTCHINSON."
736
II. SPECIAL CONTRIBUTION.
With Plates LXXV.— LXXXIII., a Map, and 15 Text-figures.
This is neither the time nor the opportunity for the composition of a costly
monograph on Nautilus. It will be time when the embryonic stages have been
discovered and opportunity when it is possible for an investigator to command the
requisite knowledge and material to permit of detailed comparisons being made, step
by step, between more or less distantly related types. I am only able to furnish
a contribution which partakes of the nature of a monograjDh in a strictly limited
sense of the term. The new matter contained in these pages consists of facts and
suggestions some of which have been already published in a preliminary form, and
relates principally to the following aspects of the subject: — bionomics (habits, range,
oviposition), branchial sense-organs (osphradia), mechanism of respiration, injection of the
vascular system, connections of the siphuncle, innervation of the ophthalmic tentacles,
development of the accessory sexual organs, enumeration of the digital tentacles,
orientation, and specific divergence.
1. Historical Survey.
The bibliography of Nautilus includes so many memoirs of the first order of
importance \mtten at varying intervals by experienced anatomists of different schools
and countries during the last seventy years, as to convey the impression that its
conchological, malacological and biological properties must be well-nigh established and
that the only scope for further contributions to knowledge must take the du-ection of
physiological and embryological researches. Such however is not the case, and so far
as the finer anatomy and cecology of Nautilus are concerned it will be long before
the Pierian spring is exhausted.
The seven decades which have elapsed since Nautilus was assigned its definite
position in the natural system fall into two very distinct periods, the first of which
was inaugurated by Su- Richard Owen^ in 1832, followed by Valenciennes^ in 1841,
and terminated by Keferstein^ in 1865. The second period, which is now in course,
1 Owen, R., " Memoir on the Pearly Nautilus." Published for the Council of the Royal College of Surgeons,
1832, with 8 plates.
- Valenciennes, A., " Nouvelles recherches sur le Nautile flamb^ {Nautilus pompilius, Lam.)." Arch. Mus.
Paris, 11. pp. 257—314, PI. viii— xi, 1841.
'■> Keferstein, W., "Beitrage zur Anatomie des Nautilus pompilius." Nachr. Ges. GiHtingen, 1865, pp. 355—
375 ; plates pubhshed in Bronn's Klassen uiid Ordimngen des Thierreichs, 1866, in illustration of Malacozoa-
Cephalopoda, Taf. ex — cxv.
HISTORICAL SURVEY. 737
was opened by the works of Professors von Jhering' (1877 — 1881), and Lankester and
Bourne = (1883).
It is a point of literary interest to note that there is some evidence, which was
discussed at length by Owen in his celebrated memoir, tending to show that the
mollusc which inhabits the shell of the Pearly Nautilus was known to the father of
natural history. However this may be, it is certain that since the revival of learning,
Nimtilus has on two occasions, separated from one another by an interval of a hundred
and twenty years, been the subject of an important zoological discovery. The first of
these was made by a Dutch physician and botanist, resident in the East Indies, named
George Everard Rumph, usually known as Rumphius, the author of a large work
entitled D'Amhoinsche Rariteitkamer, which was published in folio in Amsterdam in
1705, and was translated into German sixty j'ears later. In this volume appeared the
first recognisable descrijition and illustration of Kmitilus pompilius, this fact constituting,
I believe, one of his principal titles to enduring distinction as a zoologist.
Rumphius described some of the external characters of Nautilus with considerable
accuracy, recognising it as a Cejjhalojsod Mollusc allied to the calamaries, cuttle-fishes
and octopuses, but his statements lacked confirmation and amplification and consequently
had no marked effect on classification. Cuvier [Regne Animal, ill. p. 366, quoted by
Owen] dismissed his figure of jVautilus with the epithet " indechiffrable," but Keferstein
[Nachr. Ges. Gottingen, 1865, p. 356] was more generous, saying, " Oken allein scheint
mir diese recht gute Beschreibung und Abbildung angemessen gedeutet zu haben."
The works of Rumphius ^ ovei-shadowed b}- those of Linnaeus, became practically
forgotten, and so it happened that the capture of a unique specimen in the year 1829
by Dr George Bennett off the island of Erromanga in the New Hebrides, followed
by its successful transmission to Europe, had all the merit of an original discovery*.
In his book entitled Gatherings of a Naturalist in Australasia, Bennett tells us that
his attention had been directed to the subject of Nautilus and its relation to Ammonites
and Belemnites after a chance conversation with Professor Owen.
In the evening of August 2-ith, 1829, while at anchor in Dillon Bay, Erromanga,
he "observed an object floating upon the water, resembling a dead tortoiseshell cat.
So unexpected a sight," he says, "excited my curiosity, and the boat. ..was immediately...
sent to ascertain the nature of this floating object. It was found to be the Pearly
Nautilus On being brought on board, I observed it retract the tentacles or feelers
still closer than before; and this, with a slight quivering of the body, was the only
sign of vitality it gave." This specimen was therefore clearly in a moribund condition
' Jhering, H. von, Vergleichende Anatomie tics Nerveiisystems und Phylogenie der MoUusken, Leipzig, 1877 ;
" Ueber die Verwandtschaftsbeziehungen der Ceplialopoden." Zeitschi: wiss. Zool., Bd. xxxv. pp. 1 — 22, ISSl.
- Lankester, E. E., and Bourne, A. G., "On the existence of Spengel's olfactory organ and of paired
genital ducts in the Pearly Saiititus." Quart. J. Micr. Sc. xxiii. pp. 310—318, 4 figg. 1883. Other figures
were published in the same year in Professor Lankester's article " Mollusca," I\ncycl. Brit., '.tth ed., and
reissued in his Zoological Articles contributed to the Encyclopaedia Britanuica, 1891.
' In addition to the Rariteitkamer, he wrote the Herbarium .imboinense which was edited and published
after his death by J. Burmann, Amsterdam, 1741 — 1755.
* Bennett, G., "The inhabitant of the Pearly Nautilus." London Medical Gazette, viii. p. 729, 1831;
also by the same author, Gatheririgs of a Naturalist in Australasia; London, 1860, pp. 374 et seq.
738
EXTERNAL FORM AND PIGMENTATION,
at the time of its capture. Professor Owen, we are told, received the prize and went
to Paris to see Cuvier. But Cuvier died shortly afterwards in the same year which
witnessed the publication of Owen's memoir, without having had the opportunity of
satisfying his instinct of naturalist by the sight of this remarkable creature.
2. External Foem and Pigmentation.
The external appearance of Nautilus is too familiar to require formal description,
especially since an excellent set of drawings representing living specimens in different
attitudes has recently been published by Professor Bashford Dean'.
The complete animal consists of the shell and its tenant or, in New Britain
parlance, the " pal-a-lialia " and its " wirua," the two being inseparable until parted by
death. Although the body is firmly bound to the shell by the tension of the paired
columellar muscles, it is not really attached to the shell, the ends of the muscles being
covered over wth a horny membrane continuous with a horny girdle, so that when
the pressure of the muscles is overcome, the entire animal, including the siphuncle,
Fig. 1. Nautilus pnmpilius, L., photographed from life iu New Britain, 1895. [Reprodnced from the Quurl.
J. ilicT. Sc, Vol. xsxix. 1896, p. 179, by permission of the Editor.] About one-half natural size.
may be removed without lesion, except that the membrane is generally lacerated. The
fact that the muscles are not directly attached to the shell was pointed out by Owen
1 Dean, B. , "Notes on Living Nautilus." Amer. Natural., xxxv. pp. 819 — 837, 1901. The material was
collected in the strait between the islands of Negros and Cebu in the Philippines.
EXTERNAL FORM AND PIGMENTATION. 739
and by Vrolik', but the latter erroneously supposed that the animal was attached to
the shell by means of the siphuncle, presumably because the latter became broken off
during the removal of his specimen from the shell.
Whether at rest or on the alert, the dark brown and whitish mottled or piebald
coloration of the exposed part of the molluscous body harmonises well with the zebra-
like markings on the shell. This scheme of pigmentation apparently produces the same
effect as does the play of light on the surface ripples of the sea, and the alternating
light and dark bands of the shell might be appropriately described as ripple-markings.
On one occasion when on a raft with my native servant in Sandal Bay, Lifu,
I accidentally dropped a sprightly Nautilus overboard in four or five fathoms of clear
water. It instantly disappeared in a miraculous manner, baffling all our efforts at
recovery. Whether the gleam of colour would confer an equal degree of invisibility
in the phosphorescent twilight at greater depths it is of course impossible to say,
but when the active migi-atory habits of Nautilus are considered and also the fact
that it is subject to the attacks of voracious fishes, probably sharks and congers, as
is evidenced by the mutilated condition of many individuals, I think the above
observation may be taken to indicate that the colour-markings, being what they are,
exert a protective influence.
Under certain conditions, more frequently when kept in a confined space, I noticed
that a great portion of the brown pigment would disappear from the outer surface of
the hood, so that the latter became partly, in some cases entirely, bleached. The pigment
would seem to be actually discharged from the epidermal cells in which it is normally
lodged in an even layer near their outer ends.
The wart-like, generally whitish, gibbosities on the upper surface of the hood, have
been regarded as sensory papillae, and were likened by Van der Hoeven (1848) to the
papillae vallatae of the human tongue. It is obvious, however, that they are not
sense-organs.
In the retracted state the mouth of the shell is closed by the thick warty hood,
which forms the upper part of the cephalic comjilex and acts as an operculum. In this
condition the hood somewhat resembles the foot of a Gastropod, and Rumphius stated
that when the animal crawls the hood is directed do\vnwards, and the side which is
uppermost in the attitude of swimming becomes a jjlantar surface when creeping.
This mistake was repeated by Owen, who described the hood as being of a fibrous
texture and white colour internally, resembling dense conurii, but added that it is
" doubtless muscular, and in creeping (the position of the animal being reversed) seems
calculated to act as its chief locomotor organ." It is true that the hood is traversed .by
thin and sparse muscular fasciculi, but its structure is essentially coriaceous, and it serves
as a shield, not as an organ of locomotion.
In the attitude of swimming and breathing, as shown in the photograph here
reproduced, the hood is raised above the margin of the shell so as to expose the
tentacles, the funnel and the eyes, and to allow of fi-ee communication between the
mantle-cavity and the external aqueous medium. In the figure the front end of the
' Vrolik, W., "On the Anatomy of the Pearly Nautilus." Ann. Nat. IIUI. xii. 1843, pp. 173—175.
W. VI. 97
740 SEXUAL DIMORPHISM,
funnel may be faintly detected projecting beyond the anterior lip of the shell. The
posterior surface of the hood is concave, in accurate correspondence with the involute
convexity {anfractus) of the keel of the shell, and in the concavity there is a semilunar
fold which fits upon the blackened involute portion of the shell, towards which it acts as
a lubricant. This fold is the nuchal membrane which was compared by Keferstein (1865),
and Lankester (1883, Encyc. Brit.) with the cartilaginous nuchal plate of Sepia. I shall
have occasion to return to it later. In the specimen which I photographed, the slender
tentacles were mostly in a condition of protraction, and when, as in this instance, the
animal is stationary although expanded, they are seen to hang down listlessly, but when
swimming vigorously backwards are held out straight', like the arms of Octopus under
the same circumstances.
3. Sexual Dimorphism.
1848. Van DER Hoevex, J. Eenige afwijkingen in den vorm van het hoofd,
waargenomen bij een mannelijk voorwerp van Nautilus pompilius. Tijdschr. voor de
wis- en natuurkundige Wetenschappen, Deel i. p. 67, PI. 1, figg. 1 — 3. Amsterdam.
1850. Vax DER Hoevex, J. Contributions to the knowledge of the animal of
Nautilus pompilius. Tr. Zool. Soc. London, Vol. iv. part I. pp. 21 — 29, PI. v — viii.
1856. Van DER Hoeven, J. Bijdragen tot de ontleedkundige kennis aangaande
Nautilus pompilius L., vooral met betrekking tot het mannelijke dier. Verhandel. der
kon. Akad. Amsterdam, Deel iii., 20 pp., 5 plates.
1883. Bourxe, A. G. The differences between the males and females of the Pearly
Nautilus. Nature, xxviii. p. 580.
1895. WiLLEY, A. In the home of the Nautilus. Natural Science, Vol. vi., June,
1895, pp. 405—414.
1895. VAYSSiiiRE, A. Sur le dimorphisme sexuel des Nautiles. C. R. Ac. Paris,
June, 1895, Tome 120, pp. 1431—1434.
1901. Deax, B. Notes on living Nautilus. Amer. Natural, xxxv. pp. 819 — 837.
In Nautilus the sexes are separate, as in all other Cephalopoda, and this dioecism
is accompanied by a sexual dimorphism which is determined by the nature of the
accessory organs of reproduction and is manifested in the dimensions of the mouth of
the shell.
With regard to the statistical relations of the sexes I found that, judging from the
specimens taken in the traps, the number of males exceeds that of the females in the
case of N. pompilius, while for N. macromphalus the reverse was true. The excess of
females, or hyperpolygjTiy, is a common phenomenon among Cephalopoda, and it is
a curious fact that there should be an apparent difference in this respect between these
two species of Nautilus. The figures relating to the former species which I have
recorded are perhaps more convincing than those which concern the latter. Out of 216
specimens of N. pompilius, 150 were male and 66 female. Of 26 N. macromphalus,
1 This is well shown in some of Dr Dean's figures to which I have already referred.
SEXUAL DIMORPHISM.
741
10 were male and 16 female. The actual distribution of the numbers for N. pompilius
is given in the subjoined table.
Date
Total
Male
Female
Locality
1895.
January 18
14
12
2
Blanche Bay
22
21
16
5
»
24
8
3
5
)>
February 7
22
15
7
))
March 2
2
1
1
))
6
6
4
2
»)
7
29
22
7
»
15
10
9
1
>i
16
6
5
1
))
19
17
8
9
»
30
4
3
1
»
April 20
10
8
2
))
25
7
4
3
V
May 21
7
5
2
))
24
1
1
0
»
28
1
1
0
»>
June 13
4
2
2
»
26
6
3
3
i)
August 20
5
5
0
Talili Bay
September 13
5
5
0
J»
14
6
4
2
))
1897.
Ju]y 7
5
2
3
Blanche Bay
8
14
8
6
j»
9
6
4
2
5>
216
150
66
In contrast with the race of the Argonauts or Paper Nautili where the male is
strikingly smaller than the female, which alone carries a shell of peculiar construction,
the males of the Pearly Nautilu.i are rather larger than the females. This predominance
of the male is due to its possession of a remarkable organ, the spadix, placed beside
the mouth within the outer whorl of tentacles and formed, as first described by the
Dutch anatomist Van der Hoeven, by the combination of four modified tentacles belonging
to the inner whorl (see below). The presence of the bulky sjjadi.x in the midst of the
cephalic comple.x which, in the retracted condition, is concealed below the hood, determines
an increase in the dimensions of the latter, which must obviously find expression in the
size of the orifice of the shell.
97—2
742
SEXUAL DIMORPHISM.
The sexual dimorphism is more pronounced in Nautilus than in any other recent
Cephalopod, with the exception of Argonauta, owang to the feet that the spadix is
EXTKEME OF DlFFEBENCE.
MiNniUM OF DiFFERENXE.
Fig. 2. Secondary sexual characters in shells of Xautibis pompilius.
[Eeproduced by favour of Dr F. A. Bather from Natural Science, Vol. vi. 1895, p. 411.]
a permanent structure, having a gi-adual development from the adolescent to the adult
stages. It is not periodic in its appearance as is the hectocotylus of the Dibranchs.
That its presence should determine a difference in the shape and dimensions of the
orifice of the shell, is a fact which has already been made use of by palaeontologists,
who now recognise similar sexual variations in the shells of extinct species. Commenting
upon my observations and figures (pubUshed in 1895) relating to the shells of male and
female Nautilus, Professor Howes' pointed out that these and the nearly contemporaneous
observations of M. Vayssiere bore out the suggestion made by D'Orbignj- more than fifty
years ago that certain differences in the shells of Ammonites might possibly be indicative
of sexual variation.
1 Howes, G. B. Address of the President, P. Malac. Soc. London, Vol. n., 1896—1897, see pp. 69—71.
SPECIES AND RANGE. 743
The sexual dimorphism manifested by the shells of recent Nautilus has also been
observed among the much more numerous species which occur in the fossil state. For
example, it is shown in a striking manner in the figures of Nautilus bradfordensis Crick,
contained in an article by Mr G. C. Crick', where he described and figured the shells of
eleven species of Nautilus taken fi-om various zones in the Inferior Oolite of Dorsetshire,
of which no fewer than ten were registered as new to science.
4. Species and Range.
185.5. SowERBY, G. B. Monograph of the genus Nautilus. Thesaurus Conchyliorum,
Vol. II. pp. 463—465. PI. xcvii. and xcviii.
1855. M.\CDONALD, J. D. On the anatomy of Nautilus umhilicatus compared with
that of Nautilus poinpilius. Phil. Trans. 1855, pp. 277—288, 2 plates.
1857. Gould, A. A. On the true Nautilus umhilicatus of Lister. P. Zool. Soc.
London, xxv. 1857, pp. 20, 21.
1859. Bennett, G. Notes on the range of some species of Nautilus, on the mode
of capture, and on the use made of them as an article of food. P. Zool. Soc. London,
XXVII. 1859, pp. 226—229.
[In this paper Bennett erroneously speaks of N. macromphalus as occurring in the
New Hebrides.]
1860. Bennett, G. Gatherings of a Naturalist in Australasia. (London, J. Van
Voorst, p. 374.)
1877. Bennett, G. Notes on the Pearly Nautilus \_N. povipilius], Ann. Nat. Hist.
(4), Vol. XX. pp. 331—334.
1896. VAYSSifeRE, A. Etude sur I'organisation du Nautile (caracteres zoologiques,
dimorphisme sexuel, tentacules et spadice). Ann. Sci. Nat. (8), ii. pp. 137 — 186. PI. 16 — 19.
[Includes critical examination of the radulae of N. pompilius and N. macromphalus;
Vayssiere quotes D'Orbigny (1846, Dictionnaire universel d'hist. nat.. Article " Nautile ")
to the efi'ect that N. pompilius occurs at the Nicobar Islands "dont les habitants fument
et boucanent sa chair."]
1896. WiLLEY, A. Zoological observations in the South Pacific, in. On some
variations in the shell of Nautilus, with description of a new variety. Quart. J. Micr.
Sc, XXXIX. pp. 227—280, with figures on PL 13.
1898. WiLLEY, A. Some zoological results of a voyage to Melanesia during the
years 1894 — 1897. [N. umhilicatus.] P. Cambridge Philos. Soc, ix. p. 398.
Hitherto it has only been found really practicable to differentiate the recent species
of Nautilus according to the character of the shell in the umbilical region, and fi-om
this point of view four well-marked forms have been distinguished, namely, N. pompilius
Linn., with umbilicus concealed by a deposit of callus in the adult, N. stenomphalus
' G. C. Crick, 1898, " Descriptions of new or imperfectly known species of Nautilus from the Inferior
Oolite, preserved in the British Museum (Natural History)." P. Malac. Soc. London, Vol. in. pp. 117 — 13"J,
1898. Cf. also Buckman, S. S., and Bather, F. A., "Can the sexes in Ammonites be distinguished?"
Nat. Sci., Vol. IV., June 1894, p. 427.
744 SPECIES AND RANGE.
• Sowerby, with narrow perforate umbilicus, N. Tnacromphalus Sowerby, with rather wide
subperforate ' umbilicus, N. umhilicatns Lister, with very wide perforate umbilicus.
Judging from the known facts relative to the coiling of the shells of extinct
Ammonoids and Nautiloids and also from the fact that the young shells of N. pompilius
are perforate, the umbilical region becoming subsequently covered over by callus, I suppose
it is natural to conclude that the form of the shell in N. umbilicatus is the most primitive
among recent species, while that of j\^. pompilius is the most highly adapted.
If inquiry be made into the nature of the adaptation which has resulted in the
complete closure of the umbilicus in the latter species, it will be found as a fact that
an exposed deeply excavated umbilical region, such as is presented by the shells of
JV. macromphalus and N. umbilicatus, is liable to be selected as a nidus for the attachment
of all kinds of sedentary animals, Serpulids, Cirripedes, Ostreids, Bryozoa, Foraminifera,
and others.
The fouling of the shell resulting from the incrustations of these foreign bodies will
not only serve to initate the sensitive edge of the mantle in theii- vicinity, but ^yi\[
also materially hamper the swdmming of the Nautilus. (PI. LXXIX. fig. 1.)
The deposition of callus over the umbilicus in ^V. pompilius effectually repels the
visits of unbidden guests. In ^^e^v of these considerations it is interesting to note that
it is precisely in the umbilical region that the shell of iV. pompilius exhibits marked
variations, to which I have called attention (1896), and I came to the conclusion that
iV. stenomphalus was simply an extreme variety of this species. Shells which are assigned
to M. stenomphalus also vary in the diameter of the umbilicus, and I named a form
of which I obtained one specimen of the shell only, through the kindness of the
Hon. M. H. Moreton, at Samarai, N. pompilius, var. Moretoni, but I have since been shown
a specimen in the Manchester Museum by Mr W. E. Hoyle which is identical with mine.
In 18.55 Macdonald described the anatomy of a species from New Caledonia, which
he erroneously called N. umbilicatus, subsequently recognising his mistake, and correcting
the name to that of X. macromphalus^. Macdonald's investigation of the anatomy of
N. macromphalus resulted, as is well kno\\Ti, in the discover}- of the otocysts, but revealed
no appreciable specific differences between it and N. pomjiilius so far as the soft parts
are concerned. In fact the animals of these two species are really indistinguishable, and
I was therefore very pleased to come into the possession of a single mutilated specimen
of X. umbilicatus accompanied by its shell, which had been picked up from the surface
of the sea, not far from Milne Bay in British New Guinea, and to find that this species
differed noticeably from its congeners by the character of the hood, the gibbosities of
which have the form of flat-topped angular areas separated by deep grooves, producing
a pronounced tessellated appearance. (PI. LXXVIII. fig. 3.)
In comparison -with its former world-wide distribution, evidenced for example by the
abundance of its remains in the London Clay, the present restricted range of the genus
1 If the shell is coiled in such a manner that a space is left between the first chamber and the first
turn of the spiral so that light may be seen through the pin-hole aperture, the umbilicus is said to be
perforated. In the shell of ^V. macromphalus there is a pit on each side but it does not extend through
from one side to the other.
- Macdonald, J. D., "Further observations on the anatomy and physiology of Nautilus." P. R. Soc.
London, viii., 1857, pp. 380—382.
SPECIES AND RANGE. 745
Nautilus, shown in the map at the end of this article, appears to have a special interest
It is wholly confined to the seas adjoining the islands of the Eastern Archipelago,
including in the latter term the East Indies, Philippines, New Guinea and its dependencies,
Solomon Islands, New Caledonia, New Hebrides, and Fiji. Its shells have been picked
up on the Nicobar Islands, Japan [Dean] and the coast of New South Wales, drifting
to these localities which lie outside the range of the living animal.
It is curious that so far as is known with any certainty it does not occur west of
the Strait of Malacca nor east of Fiji. Although it descends to deep water, the single
specimen obtained during the Challenger Expedition having been dredged in 320 fathoms',
it is not an abyssal form, but rather seems to affect the vicinity of large islands, which
perhaps in former ages were united to still larger continental masses. Mr Charles Hedley,
who has developed a remarkable theory of a Melanesian plateau, which amongst other
connections united New Caledonia to New Zealand (as indicated primarily by the
distribution of land-molluscs), in a recent paper says, " It is remarkable how strictly
Nautilus observes as its eastern limit the ancient coast line of the Melanesian plateau ^"
If the generic distribution is interesting, the local range of one of the three recent
species, namely N. macromphalus, is still more strange. Although as noted above there is
no apparent difference between the animals which construct the shells of N. pompilius and
N. macromphalus respectively, yet these species are quite distinct, as is shown not only
by the coiling of the shell, which is quite constant in the latter, but also by their
geogi-aphical distribution. N. pompilius is never taken in the New Caledonian Group
[New Caledonia, Isle of Pines, Loyaltj' Islands], and N. macroinphalus is never taken
anywhere else. The species which occurs among the New Hebrides and Fiji Islands
is N. pompilius. When we consider the individual abundance of Nautilus wherever
a favourable locality is known, the circumscribed limits of the area frequented by
N. macromphalus alone are very puzzlmg. On the other hand so far as we can judge
from the scanty data which are available, the area of distribution of N. uinbilicatus overlaps
that of N. pompilius, and therefore this factor does not contribute to the differentiation of
the species, but in this case the animals are quite distinct, N. zimbilicatus, as I have
already mentioned, presenting a characteristic areolation of the upper surface of the hood,
resembling the pallial investment of Lepidoteuthis grimaldii described by M. Joubin'.
The same description applies to both sets of cutaneous structures, rhombohedral scales
of cartilaginous consistency and fibrous texture. Of course they have been independently
acquired, since they occur in different regions of the body, on the cephalopodium in
N. umbilicatus, on the mantle in Lepidoteuthis.
It follows from what has preceded that the recent species whicii has the widest
known range is N. poiupilius, since it occurs in the Philippines, Moluccas, Bismarck
Ai'chipelago, Torres Strait, New Hebrides, and Fiji. The other species which can be obtained
in abundance is confined to the New Caledonian Archipelago, namely, N. macromphalus.
Finally, the third species N. umbilicatus occurs also in Papuan waters, but the animal
I Moseley, H. N., Noles by a Naturulist on 11. M.S. ChaUeiiiifr. Edition of 1H!)U, p. 2.')(). (First edit. 187!).)
- Hedley, C, " Descriptions ol new Mollusoa, chietiy from New Caledonia." P. Linn. Soc. .V. ,5. Wale-i,
1898, Part I., see p. 100.
' Campagnes Scient. Prince de Monaco, 1900, Cephalopoda, )). 70, PI. xv. tig. 2.
746 MANTLE ; SHELL ; NUCHAL MEMBRANE.
has only been taken once. Its shell is occasionally drifted upon the shores of New
Britain and elsewhere, but it is much rarer than the others, and is often spoken of as
the "king Nautilus'^."
5. Mantle; Shell; Nuchal membrane.
Under the above heading I shall speak of the formation of the septa which results in
the incameration of the shell. The relative importance of this subject may be estimated
from the fact that there is an apparently widespread belief which requires correction.
It has been suggested, and I think in some quarters adopted as a dogma, that the
formation of successive septa is correlated with the recurrence of reproductive periods.
This is not the case since, according to my observations, propagation only takes place
after the last septum has been formed. Furthermore in spite of the numerous publications
which have dealt with the mantle, and the still more numerous treatises on the shell of
Nautilus, there is not one paper that I have been able to consult in which the delimitation
of the mantle in its relation to various parts of the shell is quite clearly displayed. The
relations of the mantle to the animal are now well known and have been so for many
years, although in one respect they were eiToneously described by Owen (1832), who has
the following statement, which must appear strange to all who have handled the animal.
He says on j). 9 of his memoir : — " At the ventral aspect of the body the mantle becomes
thinner, is prolonged anteriorly, and is perforated hy a large aperture through luhich the
funnel passes" [italics mine]. This is one of three principal mistakes- contained in Owen's
otherwise wonderful monograph, which was based upon the dissection of a single specimen,
and it is perhaps permissible to refer to them, not for the purpose of animadversion but
rather for instruction and encouragement.
The mantle is transparent in the living animal (of PI. LXXV.) and consists of two
portions, an anterior free pallial fold, which encircles the fore-part of the body like a collar ;
and a posterior portion, which forms the thin membranous integument of the visceral sac,
to which is added an appendix called the siphuncle'.
The integument of the visceral sac has always been reckoned as part of the mantle
on account of the fact that the whole of the mantle area is capable of secreting nacreous
substance externally, the posterior or \'isceral portion in particular being concerned with
the formation of the septa. The growth of the shell takes place at the free border of the
pallial fold, but the whole outer surface of the fold can deposit nacre, as is indicated by
the occasional ajjpearance of nacreous intumescences on the inner surface of the shell, and
also by the rare occurrence of the phenomenon of true pearl-formation, one example ot
which came under my observation.
' No impression of the contrast between the former and the present distribution of Nautilus as a genus
can be better obtained than by an inspection of the superb collection of Jurassic, Cretaceous and Tertiary
Nautili which is exhibited at the British Museum (Natural History). Compare also the following publications : —
Catalogue of Fossil Cephalopoda in the British Museum, Part I. (1888), by A. H. Foord ; Part II. (1891) by
A. H. Foord ; Part III. (1897), by A. H. Foord and G. C. Crick. Some seventy species of Nautilus are
enumerated in the List of the types and figured specimens of Fossil Cephalopoda in the British Museum (Natural
History) by G. C. Crick, 1898.
- The other two are the description of non-existent peripheral ganglia and the assertion of a communication
between the siphuncle and the pericardium and thence to the exterior through the viscero-pericardial apertures.
^ See below.
mantle; shell; nuchal membrane. 747
Upon taking a shell, of which the animal had died and fiillen out, from a basket
at Lifu, something rattled like a stone in the shell. It turned out to be a handsome
pearl of large size, but of doubtful value as a gem, since the shape is not quite regular,
flattened on one side, and the surface is not absolutely pure. It measures about
15 millimetres in major diameter, 11 mm. in height, weighs 3690 milligrammes and is,
I believe, the first nautiline pearl to be recorded.
The shape of the shell, a circinate spiral, belongs to one of those fundamental
forms which recur both primarily and secondarily under various guises in both kingdoms
of the organised creation. This form is calculated, for example, to meet the require-
ments of economy of space combined with strength and symmetry of construction, and
has therefore a physiological meaning. It is certainly remarkable to find a mollusc
provided with an external shell as large as that of Nautilus, which is at the same time
an expert and rapid swimmer. This faculty of swimming as the principal, practically
the sole means of locomotion, depends of course more upon the buoyancy of the shell
than upon its shape, but the latter should not be disregarded in this connection.
The buoyancy of the shell is due to the series of air-chambers which have
long excited the admiration of poets and philosophers. Successive chambers are added
by the secretion of fresh septa pari passu with the growth of the animal. If the shell,
with the live Nautilus in it, be perforated over the chambers under water, the air
bubbles gently out as the water enters. I regret that I omitted to collect any of this
air, but Vrolik (1843) ascertained that it contained more nitrogen than atmospheric air,
a fact which is not suqjrising if we consider that the animal could better afford to
part with nitrogen than with oxygen. Nautilus seems to have a peculiar faculty of
producing gas', and my knowledge of the pallial veins was chiefly due to their automatic
injection with gas after removal of the animal from the shell.
The chambers are not individually air-tight since they are perforated by the
siphuncle, but collectively they are rendered an air-tight and water-tight hydrostatic
apparatus, owing to the fact that the animal itself completely closes up the entrance
to the chambers in virtue of its adherence to the shell by the muscles and annulns
(girdle of Owen). Any loss by diffusion might be made good by the siphuncle, but
apart from this I see no reason to imagine that the air which fills the chambers
undergoes any appreciable fluctuations of pressure. It is, I am convinced, an error
to suppose that variations of pressure of the air in the chambers enable Nautilus to
rise or sink as the case may be. The air simply renders the shell buoyant once
for all.
The progressive growth of the animal is accompanied by the addition of new
chambers, which give increased buoyancy and so maintain the shell under the entire
control of the animal until a limit is reached when the last air-chamber has been
divided off, and then only does propagation commence.
' Professor Lankester (Article " Mollusea," Encyc. Brit., Vol. xvi. 9th edit.) says: — "In connexion with the
secretion of gas by the animal, besides the parallel cases ranging from the Protozoon Arcella to the Physoclistic
Fishes, from the Hydroid Siphonophora to the insect-larva Corethra, we have the identical phenomenon observed
in the closely-allied Sejiia when recently hatched."
w. VI. 98
748 MANTLE ; SHELL ; NUCHAL MEMBRANE.
There are several features in the shell which indicate that it has reached its limit
of growth and that its inhabitant is therefore mature. The lip of the shell which has
hitherto been thin and fi-agile gradually becomes thickened, and a black border is
ultimately formed round it on the inside in continuity, at the umbilicus, with the
coating of black varnish which covers the involute convexity or anfractus of the shell
from the earliest stages. The septa, as seen in sections through the shell, become
progressively thicker from the earliest to the latest, so that the last septum which
precedes sexual maturity is, when completed, the thickest of all'. Another striking
character which indicates the tennination of the chamber-building activity of Nautilus
is found in the fact that the last air-chamber of the finished shell is generally smaller
than the one which preceded it, due no doubt to advancing age, but one of my bisected
shells of N. macromphalus pro\-ides an apparent exception to this rule in that, though
the black edge has been added to the lip of the shell, the final chamber is not smaller
than the penultimate. It was formerly supposed that the most recent chamber was always,
at its first formation, smaller than its predecessor, even in young growing shells, and
this assumption was used as an argument in support of a theory of shell-growth by
intussusception [Riefstahl], but it is not so, and the error has been duly corrected'.
The total number of chambers constructed seems to vary considerably. Three shells
of JV! pompilius have 36, 34, and 33 chambers respectively, two of N. nutcromphalus have
28 and 27, one of N. uinbilicatus 32.
Although the umbilicus of N. pompilius is closed externally by a deposit of callus
it is well known that a section through the shell reveals the presence of an umbilical
fossa in consequence of which the initial chamber of the shell has a free outer surface.
In the middle of this surface when isolated by chipping awa}- the rest of the shell
there is a sub-rotund area with a slightly raised oval boss in the centre of it, and in
the middle of the boss a shallow elongate depression, the whole somewhat resembling
the structures known in plants as bordered pits. This is called after its dis-
coverer as Hyatt's scar' and is the principal feature of the shell, concerning the nature
of which and its relation to a possible protoconch we might expect to acquire special
information from a study of the embryonic stages of developments
At the median dorsal border of the earlier septa, fi-om the third to the twenty-
second or thereabouts, there is a deep pit which occasions a prominent lobe to project
backwards into the preceding chamber from each septum. This dorsal lobe of the
septum was observed by Valenciennes and later writers, and an excellent illustration has
been furnished more recently by Dr Appellcif °.
^ This condition is not shown iu many published figures of the shell, but is clearly reproduced in the
collotype published by Dr B. von Lendenfeld in his "Bemerkung zu Eiefstahl's Wachsthumstheorie der
Cephalopoden-Schalen," Zoul. Jahrb. Sijst. in., 1888, pp. 317, 318, Taf. ix.
= Bather, F. A., "The Growth of Cephalopod Shells." Geol. Mag., iv., pp. 446—449, 3 figg., 1887; also
by the same author, "Professor Blake and Shell-growth in Cephalopoda." Ann. Nat. Hist., June 1888,
pp. 421—427; also same Journal for April, 1888, pp. 298—310. E. von Lendenfeld, op. cit., 1888.
3 Hyatt, A., " Fossil Cephalopods of the Museum of Comparative ZoSlogy : Embryology." Bull. Mus.
Harvard, iii., pp. 5U— 111, PI. m. fig. 1, 1883.
* Of. Bather, F. A., "Cephalopod Beginnings." Nat. Sci., v., 1894, pp. 422—436.
° Appellof, A., "Die Schalen von Sepia, Spirula uud Nautilus." Svenska Ak. Handl. xxv., Stockholm, 1893;
see Taf. x. tig. 2.
MANTLE ; SHELL ; NUCHAL MEMBRANE. 749
The line of insertion of each septum into the wall of the shell is known as the
septal suture and follows a broadly sinuous course, well indicated for example in a
skiagraph made by Messrs J. Green and J. H. Gardiner and commented upon by Mr B. B.
Woodward'. (Cf. PI. LXXVII. fig. 1.)
In precise correspondence with the septal suture which bounds the fundus of the
animal chamber (Wohnkammer) there is an aponeurotic band round the visceral portion of
the mantle forming the boundary of the pallial area which secretes the septa (PI. LXXV.).
I figured this band in 1896, referring to it as the septal contour", and it has since
been further illustrated by Mr L. E. Griffin^ who speaks of it as the " posterior ventral
aponeurotic band," having overlooked my previous account^. Recently, however, I have
met with a still earlier description of this septal aponeurosis in the work of Appellof
(1893), though without illustration. Having referred to the annulus or homy girdle of
Owen by which the visceral sac adheres to the shell all round its circumference between
and including the shell-muscles, Appellof (up. cit. p. 75) goes on to say : — " Hinter
diesen [i.e. the annulus] und sehr deutlich markirt erstreckt sich rings um den Mantel
eine zweite Linie, welche der Septalsutur anliegt. Dieselbe tritt durch ihre wei-ssliche
Farbe scharf hervor und scheint eine Verdickung im Mantel zu bilden."
The preceding remarks lead to the consideration of the method of formation of the
septa. This is simple enough so far as the septum itself is concerned. What is not
so easy to understand is the manner in which the animal glides bodily forwards in its
shell so as to leave a space behind it which is destined to become the new air-chamber.
The foundation of a new septum consists of a very thin, easily torn membrane,
presumably conchyolin, which appears as a cuticle over the entire septal area of the
mantle, i.e. the area limited by the septal aponeurosis. Upon this membrane nacreous
matter is deposited commencing from the septal suture, and the membrane itself remains
as a thin pellicle over the hinder surface of the septum. This pellicle appears to better
advantage in some specimens than in others. It is a striking circumstance, upon removing
a young Nautilus from its shell, to find that the septum which constitutes the fundus
of the animal chamber is composed of soft membrane only without any trace of nacreous
substance. When a calcified septum is substituted for the primary membranous septum
by the deposition of nacre, it is at first exceedingly thin and ft-agile, becoming gi-adually
thickened by further secretion, but the size of the air-chamber is determined from the
commencement of calcification and undergoes no subsequent change.
If the homy membrane which covers over the muscular impressions in the shell
be removed, it will be found that each muscle-scar consists of a series of close-set
concentric lines exactly like the gi-owth-lines of the shell, not however continuous with
the latter but quite independent, superposed upon the growth-lines of the shell, and
traversing them at an angle. These superadded lines may be called the growth-lines
1 P. Malac. Soc. London, ii., 1890, p. 179, PI. xv.
•^ Willey, A., " Letters from New Guinea." Quart. ./. Micr. Sc, Vol. 39, August 189G, p. 170, fi^. Ifi (see
also below) .
" Griffin, L. E., "Notes on the Anatomy of Nautilus pompilius." Zool. Bull. (Boston), Vol. i., 1897,
p. 148, fig. 1.
^ It is indicated without a description in a drawing by .1. van dur Hoeven, Tr. Zool. Soc, iv.. Part I., 1850.
98—2
750 MANTLE ; SHELL ; NUCHAL MEMBRANE.
of the muscles or briefly the muscle-lines, in contradistinction to the growth-lines of the
shell, or simply, shell-lines. Careful inspection \vill further reveal the fact that what
I have called the muscle-lines are not confined to the actual muscle-scar, they do not
merely represent the symmetrical impressions of the muscle-fibres, but the concentric
lines are continued behind the scar as far as the septal suture, thus clearly indicating
a progressive forward movement of the muscles in correspondence with the gi'owth of the
entire animal.
It is important for a just appreciation of these details to note that the muscle-
lines on the shell are concentric with the anterior border of the muscular impression,
and have no relation to its posterior border, although the latter has a definite contour
rendered visible in spite of the continuity of the muscle-lines across and behind it, by the
fact that there is a glistening pearly sheen in the region of the muscular impression,
and merely a dull lustre behind it, though it must be added that this distinction is
more noticeable in some specimens than in others. (PI. LXXYI. fig. 9.)
The nacreous deposit which overlies the shell-lines behind the anterior border of the
muscular and annular impression, and is continued upon the face of the septum, has been
interpreted as hypostracum, the rest of the pearly substance of the shell being the ostracum,
while the external porcellanous pigmented layer is the periostracum^.
Keferstein (1865) and Appellof (1893) supposed that the mechanism of the forward
movement of the muscles in the shell consisted of a resorption of muscular substance
at the hinder border, coincident with a formation of fresh muscular substance in front.
But this pretended resorption of muscle-fibres could not be confined to the ends of the
muscles where they abut upon the shell, but must affect the entire body of the muscles.
There is no evidence whatever that anything of the kind takes place since the muscles
increase in size pari ^((6S» with the gi'owth of the animal, and the presence of the
concentric muscle-lines on the shell, visible as they are from the septal suture to the
anterior border of the muscle-scar on each side, is clearly indicative of a very gi-adual
forward gliding of the animal. As the animal grows it must of necessity move forwards
within the rigid walls of the shell, since the increase in size takes place in every dii-ection,
quite as much in girth as in length. At the same time the soft %-isceral sac can
accommodate itself to a certain extent to straitened circumstances, sufficiently to avoid
any sudden catastrophic movement, and meanwhile gas is secreted by or through the
thin septal area of the mantle, and when the limit of growth at any particular period is
reached, a new septum is laid down in the manner which I have described above. As
the animal moves forward over a tract equivalent to the interval between two successive
septa, a calcareous deposit is formed between the old and the new septal sutures.
With regard to the relation subsisting between the septal aponeurosis and the annulus,
or to keep uniformity of nomenclature, the annular aponeurosis^ which secretes the horny
girdle on its surface, it is to be noted that, on the dorsal side of the mantle these
structures are more or less confounded together, while ventrally there is in adult
1 Thiele, J., "Beitrage zur Keuutnis tier MoUusken. II. Uber die MoUuskeuschale." Zeitschr. wiss,
Zool., Bd. 55, 1893, see p. 23-i with text-figure. In the figure the author has omitted to represent the
posterior contour of the actual muscular impression.
^ Valenciennes likened the annulus to an aponeurosis.
MANTLE ; SHELL ; NUCHAL MEMBRANE. 751
specimens a wide interval between them, and in the centre of this intervening pallial
tract may be seen the line of insertion of what Huxley' named the pallio-visceral ligament,
that is, the membrane with the three fontanelles which incompletely divides the pericardial
from the visceral portion of the coelom. In young examples the interval between the
septal and annular bands is both relatively and absolutely much less considerable. These
relations also leave their impress on the shell.
The occurrence of muscle-lines on the shell- wall behind the posterior border of the
actual muscle-scar, to which I have alluded above, seems to me to be specially worthy of
note as indicating gradual normal growth. So true is this that if the horny membrane
is lost, especially in young shells, it is sometimes not easy to discern the posterior limits
of the muscular impressions, and Appellof (1893) states that in several small dried shells
of N. i^ompilius he has sought in vain for an indication of the muscle-scar and annulus.
The above description of the growth of Nautilus is in substantial accord with the
observations of Keferstein (1865), except with regard to his and Appellof's hj^aothesis
of a local resorption of muscular tissue. So far as I can ascertain, my record of the fact
that the hard nacreous calcified septum is actually preceded by a soft membranous
septum is new.
The three facts which are necessary to remember when attempting to form a subjective
conception of the mode of growth of Nautilus are as follows: —
(1) Growth in bulk of the animal taking place at regular intervals, terminated
respectively by the formation of a new septum.
(2) Rigidity of the shell necessitating a forward movement of the too bulky animal.
(3) Faculty of secreting gas.
A few words remain to be said concerning the anterior free pallial fold and the
nuchal membrane. The free border of the mantle forms a continuous collar round the
entire periphery of the animal, and may be considered in three divisions, ventro-lateral,
umbilical, and dorsal. The contour of the mantle-edge accurately coiTesponds with that
of the mouth of the shell, and the mantle is never reflected upon the external surface of
the shell, except in the region of the involute convexity (anfractus). By a coincidence
it happens that in the recent species of Nautilus the contour of the lip of the shell is
a close repetition, on an enlarged scale and in a different plane, of the shape of the
septal suture.
The dorsal fold of the mantle was described and named by Owen but not correctly
figured by him, a better illustration being furnished by Van der Hoeven in 1850 in
the Transactions of the Zoological Society. It is reflected over the involute convexity
of the shell and apparently secretes the layer of black varnish which distinguishes this
part, so that the sharp line with which this layer terminates denotes the extent to
which the mantle may be reflected over this portion of the shell. I received the
impression that the dorsal fold of the mantle does not always cover the black area but
may be retracted so that the nuchal membrane which adheres to the hinder concavity
of the hood comes to play upon this smooth surface. Frequently the animal executes
1 Huxley, T. H., " On some points in the anatomy of Nautilun pompilius." J, Liitn. Soc, iii. 1859,
pp. 3G — 44.
752 MANTLE ; SHELL ; NUCHAL MEMBKANE.
violent up-and-down movements in the shell, and at such times my observation led me
to believe that it was the nuchal membrane which plied against the black surface, and
I even thought that this membi-ane was responsible for the black deposit'. But the
evidence of sections which I have since made is against this view, and I now think
that the older writers who inferred that the black pigment was secreted by the dorsal fold
of the mantle, and Professor Joubin^ who described the gland-cells in section, are right.
The epidermal cells on the outer side of the dorsal fold of the mantle are ob\'iously
glandular and more columnar than those on the inner surface, whereas the epidermis of
the nuchal membrane is of the nature of a mucous membrane and is lower on the
side which is next to the shell than on the obverse side. I have often seen the
mantle being withdrawTi from the upper portion of the convexity of the shell when
I have been handling living N^autili, but I am unable to state the precise conditions
iinder which the dorsal fold of the mantle and the nuchal membrane respectively come
into contact with the black area. There are moreover distinct indications in the
structure of the shell itself showing that the mantle is not always under normal con-
ditions kept stretched over the entire black area, since there is another mar go limitans
situated at about one-third of the distance between the dorsal border of the septal
suture and the margo limitans of the black area. The second line to which I refer
represents the limit of a thin nacreous deposit which is laid over the black deposit on
that part of the involute convexity or anfractus (Van der Hoeven) of the shell which
faces downwards when the shell is held with the mouth directed upwards. (PI. LXXVII.
fig- 1-)
The facts may be summarised by sajang that the free lip of the shell constitutes
the margo limitans of the ventro-lateral fold of the mantle, while the anfractus presents
two margines limitantes of the dorsal pallial fold, namely, the limit of the nacreous
deposit and the limit of the black varnish. In quite small shells 33 mm. in diameter,
measured in section from the anterior lip to the convex posterior surface (the line of
maximum diameter passing some distance below the umbilicus), the nacreous deposit
extends halfway over the black area ; in older and in adult shells it occupies no more
than one-third of the black area.
The dorsal fold of the mantle has therefore two functions to perform, the deposition
of black varnish over the whole of the anfractus (involute convexity) and of nacre upon
the lower portion only.
With regard to the topographical relations of the nuchal membrane I can only
call to mind one complete and accurate account of it, namely, that given by Professor
Lankester in the article " Mollusca " {Encyc. Brit, 9th edit., 1883). Owen (1832, p. 12)
correctly describes its position and shape as a " semilunar ridge," but adds that " it is
from this ridge that the mantle is immediately continued to form the concave [dorsal]
fold." Van der Hoeven (1850) while adding a new fact repeats Owen's mistake as to
a direct connection between the nuchal membrane and the dorsal fold of the mantle in
these words : — " Under that fold [the dorsal fold] is a smaller plate of nearly the same
' Willey, A., " Letters on Nautilus, 1896." Quart. J. Micr. Sc, Vol. 39, p. 172.
- Joubin, L., " Recherches sur la coloration du tegument chez lea Cephalopodes, 4°"-' partie, Glande
s^cr^tant le vernis noir chez le NautUe." Arch. zool. exper. (2) x., 1892, pp. 319 — 324.
VENTRAL PALLIAL COMPLEX. 753
form but adherent to the posterior declivous surface of the hood and only free at its
circumference. This plate is of an aponeurotic texture and a white colour: at both
sides it is united to the dorsal fold [italics mine] and below it seems to have an
intimate connection with the two side jjarts of the funnel, and indeed to be a con-
tinuation of those parts." The new fact signalised by Van der Hoeven is the relation
of the nuchal membrane to the aloe infundibuli into which alone it passes behind and
below (PL LXXVII. fig. 4).
The umbilical portion of the pallial border is the angle which unites the dorsal
and ventro-lateral folds together, and in iV. pompilius secretes the callus which conceals
the true umbilicus of the shell.
6. Ventral Pallial Complex.
In order to render my description of the pallial topography relatively complete it
is desirable to devote a special section to an enumeration of the organs which are
contained in the spacious mantle-cavity' produced by the deep ventro-lateral fold of the
mantle. The insertion of this fold into the body-wall is not so easily understood as in
the case of the dorsal fold as it involves some of the organs, notably the branchiae,
osphradia, anus and renal organs, which in other Cephalopods fonn part of the body
proper and are not can-ied up into and upon the mantle itself.
The various organs, lines and regions which compose the ventral pallial complex
are mapped out in the coloured figure on Plate LXXV. which shows their exact normal
positions in the fresh condition as seen upon and through the thin pallial wall. Such
a figure as that referred to is serviceable from the point of view of topogi-aphical
anatomy on account of the degree of transparency possessed by the living mantle as
compared with its opacity in preserved specimens.
The mantle commences as a fi-ee fold behind the renal sacs, so that these structures
actually penetrate into the substance of the mantle, and their apertures as well as the
viscero-pericardial apertures open upon the inner surface of the mantle. The pallial
insertion of the above-named organs and also of the nidamental gland in the female,
by all of which N'autilus differs from the Dibranchs, has doubtless been noted by
anatomists who have dissected this animal, and has further received special attention from
Dr L. E. Griffin'-, who points out that "there is in the Nautilus pompilius the same
arrangement of the parts of the pallial complex as in many Gasteropoda." It may be
noted here that the particular Gasteropoda to which Nautiltis, in common with other
Cephalopoda, is specially comparable, are the Zygobranchiate Prosobranchiata which
include Haliotis and Pleurotomaria^.
' Branchial cavity of Vrolik, sub-pallial chamber of Lankester.
- GriiEn, L. E., " Notes on the Anatomy of Nautilus pompilius." Zool. Bull. (Boston), Vol. i., 1897.
Section headed "The Pallial Complex" on p. 153.
' For Haliotis see Wegmann, H., "Contributions d I'histoire naturelle des Haliotides." Arch. zool. exper.
(2) II., 1884, pp. 289—379, Pis. xv.— 3ux. For Pleurotomaria the work of the lamented Martin F. Woodward
should be consulted, " The Anatomy of Pleurotomaria beyrichii Hilg." Quart. J. Micr. Sc, Vol. 44, March 1901,
pp. 215-268.
754 SIPHUNCLE AND PALLIAL VESSELS.
In comparing Nautilus with the dibranchiate Cephalopods Professor Lankester (1883,
Encyc. Brit.) points out that " in the former the base of the fold forming the mantle-
skirt comprises in its area a part of what is unreflected visceral hump in the latter."
The consequence of this invasion of organs into the substance of the mantle is
that when the ventro-lateral fold of the latter is pulled back and turned inside out in
order to expose the pallial organs, the normal antero-posterior relations of parts are
inverted and what is actually placed anteriorly in the natural position of the mantle
(PI. LXXV.) becomes posterior when the mantle is retroverted (PI. LXXVI.). This
simple fact has led to much confusion of terminology, actual topographical conditions
being confused either with mechanically retroverted positions or with inferred moi-pho-
logical relations of parts'.
Of the nine apertures which open into or near the fundus of the mantle-cavity
only four were seen by Owen, namely the two viscero-pericardial orifices, the anus and
the opening of the female generative apparatus. Four more, the renal apertures, were
added by Valenciennes (1841), and their relations further elucidated by Vrolik-'. The
full nmnber of nine was first established in 1883 by the important discovery annoimced
by Lankester and Bourne' of the existence of a rudimentaiy antimere of the functional
generative orifice, represented on the left side by the external pore of the vestigial
structure known as the pi/riform gland.
In the male the generative pore is carried forwards and occupies a prominent
median position adherent to the veria cava, but the penis is divided by a septum
internally so that it possesses two lumina. Externally the terminal portion of the male
generative apparatus appears to be symmetrical, an appearance which is heightened
especially in the young by the existence of an arcuate radix on the left side as well
as that which leads to the so-called Needhamian vesicle on the right, but the left lumen
ends blindly behind and as shown by Lankester, Bourne, and later by Graham Kerr*,
the opening of the pyriform gland is independent and has the same relations in the
male as in the female.
In the female the genital orifice is placed on the right side precisely in the angle
made by the insertion of the mantle into the body-wall, and is represented by a
transverse bilabiate vulva which is prominent in the adult but lies deep and concealed
in the young.
7. SiPHUNCLE AND PaLLIAL VESSELS.
It may seem strange to unite these structures in one heading but it is not so strange
as it looks, especially when we remember that the siphuncle is essentially and entirely
' This point will be incidentally referred to in subsequent sections of this essay.
- Vrolik, W., (a) " Brief aan J. J. Roehussen over het ontleedkundig zamenstel van den Nautilus
pompilius." Tijdsehr. naluiirk. Wet. (Netherlands), ii. 1849, pp. 307 — 327, 2 plates. (6) "Lettre sur quelques
points de I'organisation de I'animal du Nautile flambe." Hem. Soc. Normandie, x., 1855, 16 pp., 2 plates.
^ Lankester, E. E. and Bourne, A. G., "On the existence of Spengel's olfactory organ and of paired
genital ducts in the Pearly Nautilus." Quart. J. Micr. Sc, Vol. 23, 1883, pp. 340—348.
■* Kerr, J. G., "On some points in the anatomy of Nautihts pompilius." P. Zool. Soc. London, 1895,
pp. 664—686 ; see p. 672, fig. 3 and PI. xxxix. fig. 1.
SIPHUNCLE AND PALLIAL VESSELS. 755
an appendix of the visceral sac, that is to say, of the visceral portion of the mantle,
and that it owes whatever phj'siological importance it may possess to the blood which
is supplied to it from the pallial circulation. While describing the latter we are
necessarily led to speak of the siphuncle, which is a vascular vermiform process of the
mantle and hence the reason for their association in one chapter.
By means of injections practised upon fresh specimens when I was living on the
shores of Blanche Bay, I obtained a fair knowledge of the distribution of the pallial
arteries, such in fact as had not been possible to my predecessors working ^vith pre-
served material only. I published an account of the results of my injections in 1896
{Quart. Journ. Micr. Sc, Vol. 39), which I propose to recapitulate here, the more im-
portant, in my own opinion, relating to the vascularisation of the siphuncle, which does
not, as was formerly supposed from the time of Owen's memoir (1832) to that of
Dr Haller's memoir', published in 1895, receive a main artery direct from the heart
but merely a secondary and two or three tertiary ramifications from the postenor pallial
artery.
Looked at from below through the transparent wall of the mantle the heart is seen
lying in the pericardium in the form of a transversely elongated somewhat oblong body
placed immediately behind the level of the mantle-insertion and receiving at its four
comers the paired branchio-cardiac or efferent branchial vessels (PL LXXXII.). The
hinder surface of the heart is slightly convex, while the anterior surface is emarginate,
and from the base of the groove there issues a very short vessel called by Owen the
" lesser aorta," which he correctly described as dividing almost immediately into two
branches, of which he thought the anterior was " exclusively distributed to the mucous
organ of the oviduct" (1832, p. 36), by which he meant the nidamental gland, while
the posterior passing backwards between the ovary and the gizzard entered, " without
diminution of size, the membranous tube that traverses the partitions of the shell."
A third branch (intestinal artery) arising after the bifurcation of the lesser aorta was
correctly described by Owen as being distributed to the mesentery between the ixscending
and descending limbs of the last loojo of the intestine.
The two primary branches of the lesser aorta are respectively the anterior
and posterior pallial arteries, the former supplying the anterior free mantle fold with
arterial blood, and the latter irrigating the posterior or visceral portion of the mantle,
including the siphuncle.
The anterior pallial artery bends at first inwards and downwards to the middle
line, and then runs forwards below the skin at the surface of the renal sacs. At the
point where it turns forwards a small intestinal branch (t.a.) is given off. Arrived at
the anterior limit of the region of the renal sacs, which, as already mentioned, forms
part of the free mantle-skirt, the anterior pallial artery passes into the substance of
the mantle and runs forwards towards the free margin of the latter. At some distance,
13 or 14 mm., from the free margin of the mantle, the anterior pallial artery divides
' Haller, B., " Beitrage zur Kenntniss der MorpholoRie von Nautilus pompiliua" in Semon's Zool.
Fornchungsreisen in Australien und devi Malaijischen Arcliipel., Bd. v.. Lief. ii. 1895, see p. 200, and Taf. xi.
fig. 2.
w. VI. 99
756
SIPHUNCLE AND PALLIAL VESSELS.
into two main submarginal arteries (m. a.) from which numerous radial arteries {r. a.)
proceed to the mantle-edge.
Fig. 3. N. pompilius, ? . View of ventral surface of pallial and visceral regions, to show the course of the
pallial arteries; /. funnel; m.e. free edge of mantle; r.a. radial pallial arteries; m.a. marginal pallial
artery; a.p.a. anterior pallial artery; 71.3. region of nidamental gland; b.o.a. branchio-osphradial artery;
s.m. shell-muscle; r.o. region of renal organs; h. heart; p.p.a. posterior paUial artery; g. gonad;
i.a. intestinal branch of anterior pallial artery. [From Quart. J. Micr. Sc. Vol. 39, 1896.]
Exactly at the point where the anterior pallial artery passes into the substance of
the mantle, that is to say at the level of the anterior limit of the renal region, a pair
of lateral branches arise almost at right angles to the main vessel. I suggested the name
branchio-osphradial for these arteries (b.o.a.) since among their minor ramifications they
send branches up to the tips of the branchiae suppljang the integument of the latter,
and also a small branch into each of the four osphradia. In the female they also
supply the nidamental gland. The branchial branches of these arteries are of the nature
of arterioles or intrinsic arteries of the gills in their capacity of fleshy organs.
SIPHUNCLE AND PALLIAL VESSELS.
757
The posterior pallial artery runs backwards, crossing over the ventral surface of the
heart and leaving the pericardium through the median orifice of the pallio-visceral
ligament. It then passes backwards without branching on the left side of the gonad
between the latter and the gizzard, adhering to the gonad by a ligament named by
Haller the genito-intestinal ligament. This ligament binds the gonad and intestine to
the posterior wall of the perivisceral coelom with an insertion into the wall of some
20 mm. in length running in a dorso-ventral direction. At a site near the middle
of the line of insertion of the genito-intestinal ligament into the posterior body-wall
occurs the root of the siphuncle. The posterior pallial artery reaches the posterior body-
wall through the intermediation of the free ventral border of the ligament at a point
distant by 5 mm. from the root of the siphuncle. Upon reaching, in the way described
above, what is appro.ximately the middle point of the posterior rounded surface of the
body, the posterior pallial artery passes into the integument and immediately divides
into two main branches which supply the dorsal and posterior regions of the mantle,
including the siphuncle. The two main branches of the posterior pallial artery are
the pallio-septal arteries, so called because they supply that portion of the mantle which
secretes the septa of the shell.
Fios. 4 and 5. N. pompilius, i . Two views of the dorsal surface of the pallio-visoeral region, to show the
principal ramifications of the posterior pallial artery. The siphuncular artery (a.a.) has a dextral origin
in Fig. 4, sinistral in Fig. 5.
Other letters : — m.e. free edge of the dorsal and umbilical folds of the mantle ; c. region of crop ; s.p. siphuncle ;
I. region of liver; p. p. a. posterior pallial artery; yiz. region of stomach; i/. region of gonad; i. region of
intestine. [From Quart. J. Micr. Sc. Vol. 39, 1896.]
A variable number of small arteries are distributed to the siphuncle, but there is
one branch, a secondary branch of the posterior pallial artery, which may be regarded
99—2
758 SIPHUNCLE AND PALLIAL VESSELS.
essentially as the siphuncular artery. This arises indifferently ti'om either the right or
the left of the two principal branches. Owen (1832), Valenciennes (1841), Vrolik (1855),
and Haller (1895) have described what I have called the posterior pallial artery as
passing direct to the siphuncle. Keferstein (1865) with more accuracy said that it
passes " nach hinten zur Korperhaut und besonders zum Sipho." Injections show that
the sijihuncular artery is simjily on a par \vith the other ramifications of the posterior
pallial artery, the ultimate branches of which constitute a remarkably rich system of
irrigation for the septum-producing area of the mantle.
Before continuing my account of the pallial arteries it will be convenient at this
point to complete what I have to say regarding the connections of the siphuncle.
Owen, as I have already mentioned above, described a direct communication between
the pericardial cavity and the siphuncle, sajdng (memoir, p. 27): — "by the side of this
vessel " [the posterior pallial artery] " a free passage is continued between the gizzard
and ovary into the membranous tube or siphon that traverses the di\'isions of the shell ;
thus establishing a communication between the interior of that tube and the exterior of
the animal " [through the viscero-pericardial apertures].
Valenciennes (1841) denied any communication between the siphuncle and the
exterior through the mediation of the pericardium, and further affirmed that it did not
even open into the abdominal ca\'ity at its base. Vrolik (1855, p. 7) said that
Valenciemies was wrong, " car, d'apres ce que j'ai vu, le siphon s'ouvre dans la ca\"ite
abdominale, comme la pi. i., fig. 5 c le demontre." But on turning to the figure referred
to we find that it is by no means demonstrative in this respect. Vrolik was confident
as to the actuality of his observations, and stated in a foot-note that he had shown
his preparation to many foreign anatomists, including Kolliker, who mentioned it among
the impressions of his travels in his Zeitschrift. It is no light matter to attribute an
error to an anatomist of Vrolik's reputation, especially since he alludes at another part
of the " Lettre " fi"om which I am quoting to " I'erreur grossiere de voir des ouvertures
la ou elles ne sont pas." Nevertheless I shall proceed to show that he was wrong in
this matter in spite of the asseverations of Keferstein (1865) and Haller (1895).
Keferstein' defines the siphuncle in these words : — " Der Sipho ist hinten eine rohrige
Fortsetzung der Korperhohle, ^vii-d der ganzen Lange nach von einer Arterie durchlaufen
und muss grade wie die Intervisceralraume des Korpersackes venoses Blut enthalten
konnen."
It is true that in addition to its main artery, the siphuncle is travereed by a wide
axial cavity, and it is perhaps to be inferred from Keferstein's definition that this axial
ca\'ity of the siphuncle is a continuation of the abdominal or peri\'isceral ca\-ity in
which the gonad and other viscera lie, in other words that it is part of the secondary^
body-cavity or coelom. But whatever inference may be dra^vn from somewhat vague
statements, we find precise assurances on this subject in the recent memoir of Dr Haller
(1895, p. 201), who says : — " Der Sipho communicirt direct mit dem grossen Colomraum,
und zwar, da sich liber ihrer Miindung der ventrale Theil des Genitointestinal-ligamentes
an die hintere Wand des Coloms befestigt, scheinbar mit zwei Oeffnungen, thatsachlich
1 Sachr. Ges. Gottingen 1865, p. 369.
SIPHUNCLE AND PALLIAL VESSELS. 759
aber nur mit einer. Somit ist das Innere vom Sipho als die directe Fortsetzimg des
grossen Colomraumes aufzufassen, und das Colomepithel setzt sich als ein niedrig-
cubisches, beinahe plattes Epithel in die Siphonalhohlung continuirlich fort."
The relative importance of a correct knowledge of the nature of the siphuncular
cavity is too obvious to render an apology for the quotations which I have given
necessary. In fact this cavity is a venous cavity or haemocoel, as I have satisfied
myself by means of transverse and longitudinal sections, but above all by dissection,
which besides showing the non-existence of any opening into the coelom has revealed
the presence of a hitherto undescribed venous channel, which I will call the posterior
pallial vein, into which the siphuncular vein, i.e. the axial cavity of the siphuncle,
discharges (PI. LXXVI., figs. 7 and 8). It is quite probable that the posterior pallial
vein which runs above, approximated to, and parallel with, the posterior pallial artery
in the genito-intestinal ligament, is identical with the " free passage " which Owen
described as coursing alongside of the posterior pallial artery between the pericardium
and the siphuncle. As a matter of fact the chaimel which occurs in this position is
a vein which runs forwards until it arrives at the pericardium, when it bends up above
the dorsal wall of the latter and enters the central venous sinus from whence the
afferent branchial vessels are given off.
The finer structure of the siphuncle (PI. LXXXIII.) is complicated and not readily
intelligible. Surrounding the central vein is a loose meshwork of trabeculae, the meshes
communicating with one another and with the central vein. The communications
between the peripheral system of spaces and the central sinus are best seen in longi-
tudinal sections, but are also to be found in transverse sections. The external epithelium
of the true or pallial siphuncle which secretes the partially calcified and partially horny
siphuncle, which remains behind after the soft parts are extracted, presents a very
peculiar structure. This has been in part correctly described by Haller, who states
{op. cit., p. 201) that the entire surface of the siphuncle is thro\vn into very fine
longitudinal folds due to the plaited membrana basilaris of the epithelium, and in each
plication of the basement-membrane there is a narrow space lined by flattened endo-
thelium (Haller, op. cit, Taf Xii., fig. 19).
I can confirm the existence of these folds, although they do not appear in my
preparations to be so regular as Dr Haller describes and figures them. Moreover I cannot
see the regular deep epidermal pits which alternate in correspondence with the subjacent
mesodermal folds in his figures. What seems very remarkable is the fact that the
spaces in the folds are in reality prolongations from the meshes of the trabecular tissue
of the siphuncle, and therefore represent intra-epidermal blood-spaces. I think this
circumstance must throw some light on the function of the siphuncle, which is essentially
a vascular appendix. I have already expre.ssed the po.ssibility that the siphuncle would
be capable of keeping the air in the chambers up to the normal pressure by making
good losses caused by diffusion. The same, or a closely similar suggestion, was in fact
made by Keferstein (1865, op. cit. p. 374)'.
Coupling the existence of intra-epidermal venous spaces with my observation recorded
' Cf. Meigen, W., " Ueber den hydrostatisehen Apparat des Nautilus pompilius." Arch. Natury. 3G Jlig.
Bd. I. 1870, pp. 1 — 34. This author confirms Keferstein's theory.
760 SIPHUNCLE AND PALLIAL VESSELS.
above, of the automatic injection of the pallial veins with gas (PI. LXXXII.), it seems
that the conclusion may be drawn that the separation of gas rich in nitrogen is a special
property of the venous system of Nautilus. Whether this supposition will prove to be
sound or not, the observations are sufficiently singular to merit the attention of future
investigators.
The sj3ong)' trabecular tissue of the siphuncle becomes looser, i.e. the meshes become
larger near the root of the siphuncle, and where the latter abuts upon the body-wall
there is generally to be seen from the inside of the perivisceral coelom a large cushion,
divided superficially into two equal or nearly equal portions, by the insertion of the
genito-intestinal ligament. In certain conditions of preservation the wall of this sac
is very easily ruptured, so as to produce the appearance of a natural orifice, although
such is not and cannot be present, since the cavity of the sac is the radical sinus of
the siphuncular vein, and opens directly into the posterior pallial vein. When the sac is
cut open it is found to contain a fiocculent substance, composed partly of blood, and
partly of the trabeculae described above.
The radical sinus of the siphuncle of Nautilus is obviously identical in principle
with the pallio-siphonal sinus described by Huxley and Pelseneer' in their joint work
on Spirula. In Spirula we are told that the cavity of the membranous siphuncle is
a blood sinus which opens at the root of the siphuncle into a relatively large space,
which is the pallio-siphonal sinus. The irrefutable demonstration of the existence of
corresponding connections in Nautilus seems to me to be a matter of uncommon interest.
The peculiar segmentation of the siphimcle due to shallow constrictions at the points
where it traverses the septa was first observed by Valenciennes. According to Appellof
{op. cit. 1893, p. 79 et seq.), it is open to question how far back the living siphuncle
extends in the adult shell, and it is even quite certain that it does not extend into
the fii-st chamber, since he has been able to confirm Hyatt's statement "dass auch der
Sifo der zweiten Kammer wirklich blind geschlossen ist." In extracting the
animal from the shell, vaiying lengths of the siphuncle emerge. I cannot assert that I
have ever obtained a complete siphuncle, but often one with upwards of twenty segments.
I made some experimental observations on Nautili with severed siphuncle for the
purpose of ascertaining whether this organ performed a definite immediate function. The
result, so far as it went, seemed to indicate that the cutting of the siphuncle does not
temporarily affect the vitality of the animal, does not inhibit movements of translation
and does not prevent it from floating at the surface, nor from sinking to the bottom.
Its function must therefore be a cumulative one, a conclusion in accord with the
suggestion that it may serve to keep the air-chambers at the proper tension. I found
that the best way of performing the operation was to saw through the shell in the
neighbourhood of the posterior portion of the body of the animal over the cardiac region.
If the shell with the li\'ing animal inside be held mouth downwards, the cardiac region
lies approximately in the same vertical transverse plane with the points at which the
' Huxley, Rt. Hon. T. H. , and Pelseneer, P., " Report on the specimen of the genus Spirula collected
by H.M.S. Challenger." Published as the 83rd and last part of the Zoological Series of Challenger Reports,
and inserted as an Appendix to the Summary of Results, second part, 1895. Plates by Huxley, text by
Pelseneer. For the account of the pallio-siphonal sinus see p. 24, text-fig. Q.
SIPHUNCLE AND PALLIAL VESSELS.
761
free lip of the shell merges into the umbilicus. When a large enough hole has been
made in the shell to admit the scissors, the shell being still held upside down, the
postero-ventral Wsceral portion of the body behind the annulus usually detaches itself
from the shell, or can be readily caused to do so, and sinking inwards, exposes the
root of the siphuncle, which can then be .severed. Upon righting the shell the body
resumes its normal contact with the fundus of the shell, thus preventing an immediate
and extensive hemorrhage. For example a Nautilus treated in this way, and replaced
in the sea at a suitable spot, swam about vigorously for some time in the middle stratum
m.fuz
p.s.a. -I
post.p.a.
Fig. 6. iV. pompilius, i . Dorso-posterior aspect of visceral region, to illustrate the circulus pallialis and the
septal contour; n.m. nuchal membrane; n.a. nuchal artery; m.e. free mantle-edge; p.n.a. pallio-nuchal
artery ; m.p.a. marginal pallial artery ; coL columellar or shell muscle ; p.c.a. posterior columellar artery ;
p.p.a. posterior proveutricular artery ; s.c. septal contour ; s. siphuncle ; p.s.a. pallio-septal arteries ;
s.a. siphuncular artery; /. liver; i. intestine; t. testis; post.p.a. posterior pallial artery; ff. gizzard.
N.B. — The dorsal aorta and its branches are indicated by dotted lines. They show dimly through the skin
when injected. [From <^«iir(. ./. Micr. Sc. Vol. 39, 1896.]
of water, but chiefly at a little distance from the bottom. Others remained floating and
swimming about on the surface during the whole time of observation. They did not
762
SIPHUNCLE AND PALLIAL VESSELS.
go far in one direction, but tended to move in circles, as they sometimes do without
severance of the siphuncle. If one of the individuals floating at the surface was forced
down to the bottom with a hand-net it would slowly rise to the surface again, as also
often happens with a ±i^autilus which has not been operated upon. Another specimen
showed the opposite tendency to sink to the bottom very gradually.
Fio. 7. iV. pompilius, i . View of nuchal region, to farther illustrate the circulns paJlialis. The dorsal free
mantle-edge is reflected and a median incision made. h. hood ; con, concavity at base of hood, in which
the nuchal membrane (n.m.) lies; /. dorso-posterior portion of funnel; n-a. nuchal artery; col. columellar
muscle; m.p.a. marginal pallial artery; p.n.a. pallio-nuchal arteries; d.a. dorsal aorta; m.e. free mantle
edge; c.e. cut edges of mantle and body- wall.
N.B. — The dorsal aorta shapes its course in this region in accordance with the state of repletion of the crop.
[From Quart. J. ilicr. Sc. Vol. 39, 1896.]
Some further features in the pallial circulation of Nautilus still remain to be
described before concluding this chapter. After successful injections a striking peculiarity
FUNNEL AND CAPITO-PEDAL CARTILAGE. 763
in connection with the anterior pallial artery may be observed. It will be remembered
that upon approaching the mantle border in the mid-ventral line, this artery bifurcates
into two submarginal arteries (m.p.a.).
The submarginal pallial artery is continued dorsally on each side into a branch of
the systemic or greater aorta, so that a complete arterial circuit, the circulus palliulis,
is achieved. I have even partially injected the pallial arteries from the dorsal aorta
itself, although naturally the fluid did not proceed very far in the centripetal direction.
Thus by means of the submarginal arteries the system of the lesser aorta becomes
confluent with that of the greater aorta, or in other words, there is an anastomosis
between the pallial circulation and the systemic circulation.
In one of the figures here reproduced the union of the submarginal arteries with
the dorsal aorta is represented as seen through the mantle ; in the other it is sho\vn
after the mantle has been slit open and turned back, and a further incision made into
the body-wall in the nuchal region. The vessel which eifects the junction is called
the pallio-nuch'il artery {p.n.a.). The actual point where the confluence takes place
corresponds with the umbilical region of the mantle, and with the angle of insertion
of the mantle in this region into the body-wall.
From the angles formed by the confluence of the pallio-nuchal with the submarginal
arteries a branch is distributed forwards to the nuchal membrane. (Fig. 7.)
Finally I have to record some observations in the pallial veins. Under ordinary
circumstances one might never suspect the occurrence of definite venous channels ia
the mantle. When a Nautilus kept in a confined space becomes moribund, it usually
rises to the surface, owing apparently to an abundant production of gas in the interior of
the body. If it be allowed to die and be then removed from the shell the veins are
found to be injected with gas, most likely due to a regurgitation of gas, if the expression
may be allowed, through the siphuncular venous sinus and posterior pallial vein, into
the general venous or lacunar system of the body. Under these conditions the finest
ramifications of the veins, especially in the free mantle-flap, are displayed with a clearness
which could hardly be attained, certainly not surpassed by artificial injection. The mantle
in fact is seen to be riddled by these veins, which are collected into two main channels,
placed on either side of the anterior pallial artery, and discharging behind into the
afferent branchial vessels. At the sides of the mantle there may be seen a number of
lateral pallial veins, which appear to open into a large sinus situated over the shell
muscle. (PI. LXXXII.)
8. Funnel and Capito-pedal Cartilage.
It is a fact of some interest which deserves to be brought into relief, that the
cartilaginous endoskeleton of Nautilus essentially belongs to the funnel, it is the
sustentaculum infundibuli besides serving incidentally as the foundation and support of
the entire cephalopodium, and affording a fulcrum for the insertion of all the principal
muscles of the body. It agrees analogically in point of form with the hyoid cartilage
of Vertebrates and the nuchal chondroid skeleton of Enteropneusta, consisting as it does
w. VI. 100
764 FUNNEL AND CAPITO-PEDAL CARTILAGE.
of a median body with anterior and posterior comua, the whole being shaped somewhat
like the letter H. It has frequently been designated by the same term which is applied
to the principal cartilage of the Dibranchs, namely, cephalic cartilage\ but this name
as applied to the cartilage of the Tetrabranchs is not only a misnomer but is positively
misleading, since the cephalic cartilage of the former is a cranial cartilage having essential
topographical relations with the central nervous system, and presenting definite fossae for
the reception of the cerebral, pedal and \4sceral ganglionic centres, and for the protection
of the otocysts, and foramina for the exits of nerves.
In Nautilus the arrangement is more primitive in so far that the cartilage has essential
relations with the motor system generally (funnel and musculature), and merely exhibits
incidental contiguity with regard to the more ventrally situated parts of the nervous and
sensory systems. It surrounds no part of the last-named systems, is not perforated by
any nerves, and is only traversed by one pair of blood-vessels, namely, the infundibular
arteries (PI. LXXXI.). The principal fossae which it presents are related to parts of
the venous system, e.g. there is a deep fossa on each side nearly enclosed by cartilage,
for the passage of the infundibular vein, and a spacious median fossa for the anterior
reservoir of the vena cava. The deep fossa described by Owen for the lodgment of the
optic ganglia has no existence.
The anterior cornua of the cartilage of Nautilus extend forwards into the substance
of the funneP, which they help to keep open just as the posterior cornua of the skeleton
in Enteropneusta keep the mouth open ; the posterior cornua occupy the raphe which
separates the siphonopodium (funnel) from the cephalopodium; finally the body of the
skeleton lies at or near the angle of insertion of the funnel into the body-wall.
The ventral surface of the body of the cartilage is produced into a median keel
which serves for the insertion of the transverse muscles at the base of the funnel, and
its upper surface is complicated by its special relation to the important venous sinus,
which collects the blood from the head and funnel and passes it on to the vena cava.
This sinus may conveniently be termed the endochondral sinus'^.
After what has been said it will be evident that the term " capito-pedal " introduced
by Professor Lankester*, describes the cartilage of Nautilus more appropriately than any
other available expression. Nevertheless the cartilage undoubtedly affords some support
and protection to the ganglionic complex and otocyst, especially the latter, since conjunctive
trabeculae traverse the space between the perineurium and the perichondrium, and also
between the periotic membrane and the latter, in fact the loculus in which the otocyst
lies is bounded posteriorly by the perichondrium and anteriorly by the brachial or pedal
perineurium.
The dfrection of the long axis of the cartilaginous skeleton of Nautilus is probably
of importance in regard to the more general question of Cephalopod orientation, and it
' " Kopf knorpel " of Keferstein 1865 and Haller 1895.
- "Within the crura of the funnel" (Owen).
3 The endochondral sinus was described by Owen in the following words (memoir, p. 16): — "In the body
of the skeleton is excavated a large sinus, which receives the contents of the veins of the head and funnel,
and empties itself into the commencement of the great dorsal vein." The "great dorsal vein" of course
refers to the I'ena cava which is quite superficial and ventral, as Owen correctly states on p. 27 of his memoir.
* Art. "MoUusca," Eiu-yc. Brit. 9th edit. 1883, Zool. Articles (1891), p. 143.
FUNNEL AND CAPITO- PEDAL CARTILAGE. 765
may be gathered from such a dissection as that represented in fig. 1, PI. LXXXL, that
the main axis of the cephalopodium. as indicated by the general trend of the tentacular
sheaths, does not coincide with the skeletal axis but describes an angle with it. I shall
refer to this matter again under the heading " Flexure and Orientation," but meanwhile
I cannot help thinking that the occurrence of conflicting axes in relation to the
cephalopodium of A\iutilus may have morphological significance, as affording evidence
of profound topographical changes which have upset the primitive axial simplicity. Such
an axial disturbance is no longer apparent in the Dibranchiata, so that whatever it may
indicate, it is at all events a remarkable feature in the organisation of Nautilus and
should be, though it never has been, taken into account.
The limit of the extension of the anterior comua into the crura of the funnel is
superficially indicated on each side by the presence of a limbate border (l). These comua
do not reach to the base of the tongue-like valve which is inserted upon the inner
surface of the dorsal wall of the funnel.
The posterior comua underlie the optic ganglia, the olfactory labjTinths (rhinophores)
and the otocysts in the order named (cf PI. LXXXI. fig. 5), but do not embrace these
structures. Into the hinder extremities of the posterior comua are inserted the retractor
muscles of the buccal cone (PI. LXXXI. fig. 7), and the angles in which the insertion
takes place constitute deep pits in the fundus of the peristomial haemocoel.
The funnel itself comprises several regions which differ in colour, texture and function.
Its anterior and posterior portions are separated from one another by the anterior cornua
and body of the cartilage. At the forward limit of the latter there occurs externally
a peculiar border or liinbus, the linibus in/undibuli (1). If the funnel be cut across behind
the livibus the whitish crura in/undibuli penetrated by a pair of subcylindrical muscles,
the levatores in/undibuli of Owen, lying in their own venous sinus on each side, are
seen to be clearly separated by the anterior cornua of the cartilage, from the soft brownish
yellow alae in/undibuli which extend backwards and dorsally to the base of the nuchal
membrane. (PI. LXXXII. fig. 3.) It is one of the best kno\vn peculiarities of the
organisation of Nautilus that the funnel is cleft down the ventral surface, so that its
flaps overlap and present a free border throughout their entire length. The anterior
portion of the free borders of the flaps belongs to the region of the cruia, and the
posterior portion to that of the alae, although of course quite continuous. The crura
are united together dorsally by the thin median wall of the funnel which rests in the
infimdibular fossa of the great oral sheath and carries, on its inner surface, the large
valve of the funnel. Behind this thin-walled anterior portion, the dorsal wall of the
funnel, between the alae, is greatly thickened owing to the presence of the spongy
compressor muscle, the transverse fibres of which are inserted into the median keel of the
cartilage {carina cartilaginis).
In the fresh condition the integument which covers the compressor muscles and the
adjoining surface of the ala£ in/undibidi is found to be a slimy, mucous membrane, forming
two large pads meeting in the middle line and constituting the mucus-gland of the funnel.
It is interesting to note that Jatta' has interpreted the funnel-organ of Dibranchiate
' .latta, G., "Sopra I'organo dell' imbtito nei Cefalopodi." Boll. Soc. Xtrpoli (Ser. 1) vii. 189.S, pp. 4.5—60,
Tav. IV. The funnel-organ is eometimes called the "Organ of Verrill" or "Organ of Miiller." Cf. Hojle, W. E.,
" ObBervations on the anatomy of a rare Cephalopod (Gonatus fabricii)." P. Zool. Soc. London, 1889, p. 128.
100—2
766 FUNNEL AND CAPITO-PEDAL CARTILAGE.
Cephalopoda as a mucus-gland, and has further homologised it with the pedal gland of the
Gastropoda.
After comparing the figures published by Jatta, relating to the funnel-organ of
Sepia offldnalis, Octopus vulgaris and other species, I have no doubt that the funnel-
gland of Nautilus is an organ homologous with the coiTesponding structure in the funnel
of other Cephalopoda. I have not made sections through this gland, and shall have no
further opportunity of doing so, but I have really no hesitation in the matter notwith-
standing this omission (cf. PL LXXVI. fig. 1).
The funnel-organ of Nautilus passes laterally into the alae infundihuli. and posteriorly
it is attached to the ventral surface of the skin covering the shell-muscles by an elongated
limhus. I observed a network of what I took to be venous channels below the viucosa
of the funnel-organ, and two large veins passing from the limbus into the ve)ia cava.
(See figures on PL LXXXII. and LXXXIII.)
The cartilage forms a very incomplete wall to the endochondral sinus, only the ventral
and lateral sides being bounded by massive cartilage, while dorsally there is a narrow bar
of cartilage (a7xus cartilaginis) which forms two bosses projecting into the peristomial
haemocoel between the pedal and visceral commissures. Between the arcus and the pedal
commissure are to be seen two orifices which represent main conduits leading fi-om the
peristomial haemocoel to the endochondral sinus (PL LXXXII.). Behind the arcus occurs
the insertion of the inferior retractor muscles of the buccal cone into the body of the
cartilage on each side.
The infundibular nerves arise fi-om the pedal commissure and run straight forwards
into the crura infundihuli dorsad and ectad of the tubular sinuses, in which the above-
mentioned levator muscles lie. Their topogi-aphy will be elucidated in the chapter on
the peristomial haemocoel.
In concluding the present cha23ter, I should like to emphasize the \-iew that the
capito-pedal cartilage of Nautilus is to a large extent a funnel-cartilage, and is probably
not directly homologous with the cej)halic cartilage of Dibranchs, but related to the
latter through substitution. It is of coui-se obvious that the principle of substitution is
apt to become little more than a form of words, and to convey no true image to the mind.
In regard to the endoskeleton of Cephalopoda the idea which is present to my mind is
the following. The cephalic cartilage is to the Dibranchs what the capito-pedal cartilage
is to Nautilus, but the former i.s not to be identified with the latter, and does not
merely represent a specialisation of it. If this be so we might expect to find traces
of a true capito-pedal or funnel-cartilage in the Dibranchs; and indeed a pair of siphonal
cartilages' curiously recalling the cornua of the cartilage of Nautilus, has been discovered
by Hoyle- in the remarkable and, in more than one sense, primitive genus Gonatus.
Without venturing to express the definite opinion that these apparently rudimentary
siphonal cartilages of Gonatus ma}' represent vestiges of a capito-pedal cartilage such as
occurs in the still more primitive genus Nautilus, I think the suggestion may be one
worthy of the attention of teuthologists, although the fact that they occur in the ventral
' The funnel of Cephalopoda is often quite properly called the siphon but the latter term is also quite
improperly applied to the siphuncle of Tetrabranchiates, and for this reason I have avoided its use.
- Hoyle, W. E., "Observations on Gonatus fabricii," op. cit., see p. 120. Cf. also on Gomitus, the
same author's Report on the Cephalopoda collected by H.II.S. Challenger, 1S86, p. 173.
DIGITAL AND OPHTHALMIC TENTACLES. 767
wall of the siphon of Gonatus might be thought to militate against this view. Still
the anterior comua of the cartilage of Nautilus penetrate into the ventral wall of the
funnel.
9. Digital and Ophthalmic Tentacles.
The head or cephalopoclium of Nautilus comprises the better half of the animal, and
represents a complication of organs and appendages performing divers functions. The
body-wall in this region is composed in the maia of a thick coriaceous integument, which
presents a dorsal convexity formed by the mottled bro\vn-coloured hood, and three deep
concavities, anterior, posterior and ventral. The anterior deepest concavity is the fossa
huccalis, which is surrounded by the tentacular crown and lodges the great buccal cone;
the posterior conca\'ity is the fossa nuchalis which receives the anfractus of the shell,
and gives origin to a muscular Hap, the semilunar nuchal membrane ; finallv the ventral
concavity is the fossa infundibularis, which varies somewhat in depth according to the
state of muscular contraction of the parts concerned in its formation, and is exactly
suited to receive the funnel.
The cephalopodium of Nautilus consists of a tentacular crown disposed about the
mouth in the form of an oral sheath, and composed of two whorls, outer and inner.
The outer circlet' of tentacles comprises the " digital processes " of Owen, while the inner
circlet consists of the so-called " labial processes," but the latter is an unfortunate term,
since the actual buccal orifice is provided with a prominent lip surrounding the mandibles
and richly furnished with labial papillae. I therefore propose to refer to the tentacles
generally, whether belonging to the outer or to the inner whorl, as digital tentacles. The
two tentacles, the one in front of, and the other behind the eye on each side, which
Owen named " ophthalmic tentacles," really belong to the outer whorl, but have become
so modified in function and position as to require separate treatment.
The tentacles arise from the bottom of profound sheaths, ^vithin which those of the
outer whorl are completel}' retractile and those of the inner whorl partially so, but in
either case it is at once apparent that the tentacles are essentially marginal appendages.
The digital tentacles have been more particularly described as to their structure
and distribution by Owen (18.32), Valenciennes (1841), Owen^ (1843), Lankester and
Bourne (1883), Vayssiere (1896), Willey^* (1897), Griffin'' (1898). These tentacles have
in general a trihedral form tapering slightly towards the tip, and are divided by shallow
annular grooves into a large number' of articulations, the grooves being deeper on the
' The outer whorl is the same as Lankester's " auuular lobe" (Article " MoUusca," Encyc. Brit. 9th Edit.).
- Owen, E., "On the structure and homology of the cephalic tentacles in the Pearly Snutilus." Ann.
Nat. Hist. xn. 1843, pp. 305—311, 2 figg.
^ Willey, A., " The pre-ocular and post-ocular tentacles and osphradia of Xtiutilus." Quart. J. Micr. Sc.
Vol. 40, April 1897, p. 197; also in same volume (Aug. 1897), "The adhesive tentacles of Xautilm "
p. 207.
* Griffin, L. E., " Notes on the tentacles of Nautilus potnpilius." John-i Ilupkiiu U/iiv. Circ. xviii.
Nov. 1898, p. 11; reprinted in Ann. Nat. Hist. (7) m. Feb. 1899, p. 170.
' Sixty to one hundred, according to Jlr Griffin.
768 DIGITAL AND OPHTHALMIC TENTACLES.
inner sides which form the bases of the trihedra, so that a series of transverse ridges
is produced, by means of which the tentacles are able to adhere to surfaces in a manner
analogous to the action of the feet of a gecko. When attached to a rough surface by
means of the suctorial ridges the tentacles may be drawn out to a great length in all
directions, and hold on to their support with so much tenacity that sometimes when
I forcibly detached the animal some of the tentacles broke off and remained fixed to
the substratum. The suctorial ridges of the tentacles have a pale neutral tint in the
fresh state, while the crests and sides of the trihedra are distinguished by a deep
browTi pigment. The tentacles which thus act in a way resembling the arms of the
Dibranchiate Cephalopoda are the digital tentacles of the outer whorl. They have
an adhesive, prehensile and tactile function.
In the middle of each tentacle runs a large axial nerve, which originates directly
from the ganglionic nerve-centre and presents, as was first noted by Griffin, a series of
slight ganglionic enlargements at its periphery, in correspondence with the annulation
of the tentacle. In sagittal section the axial nerve is seen ^vith its peripheral ganglion
cells which constitute an almost unbroken layer on the ventral side of the nerve, that
is on the side nearest to the suctorial ridges, but on the opposite side are segi-egated
into groups. The suctorial ridges appear as bold outstanding nearly flat-topped excrescences
slightly convex, with a tendency to imbricate at their edges. They are covered with a high
and very dense epidermis which passes abruptly into the indifferent layer which lines
the intervening grooves. Below the nerve the tentacle is traversed by an artery, and
below that again, a venous channel'. Owen (1843) gi'aphically described the dense central
tissue which protects the nerve and artery, fi-om whence radiating fasciculi proceed to
the periphery ^vith longitudinal muscular bundles in the meshes. He also pointed out
how similar requirements elsewhere are met by analogous structure. Thus " the mechanical
arrangement of the contractile fibres is very similar to that of the complex muscles
described by Cuvier in the proboscis of the elephant. The attenuation and elongation
of this brobdignagian [sic] tentacle must be effected without compressing the central
breathing-tubes and the transverse fibres accordingly radiate from the dense ligamentous
tissue which suiTounds the tubes." We may add that a closely analogous structure is
met with in the proboscis of the Enteropneusta. The arms of the other Cephalopoda
are also built upon the same plan. Owen duly noted the suctorial ridges which he
described as presenting " the appearance of a number of close-set transverse plates." He
says further that " this modification must increase the prehensile and sentient properties
of the inner surface of the tentacle, and it is on the corresponding part of the larger
and fewer tentacles of the Dibranchiate Cephalopods that the acetabula are developed."
Again on p. 310 (1843) he states that "the prehensile surface of the tentacula of the
Nautilus is made adhesive, after the type of the simple laminated sucker of the Remora."
Finally the primary import of the tentacles of Nautilus is " plainly indicated by the
direct derivation of their central nerve from the cephalic ganglion."
These quotations serve to show that Owen had acquired a very just appreciation
of the general nature of the digital tentacles, although it should be noted that the fact
' A good representation of a transverse section through a tentacle is given by Vayssi^re in Ann. Set.
Nat. (8) n. 1896, PI. 19, fig. 19.
DIGITAL AND OPHTHALMIC TENTACLES. 7G9
of adhesion by means of what I have called the suctorial ridges had not then been
established.
Owen's paper of 1843 was written by way of reply to Valenciennes (1841), who had
compared the sheathed tentacles of Nautilus not with the arms of the Dibranchs, but
with the suckei-s which are borne upon the arms, the sheath corresponding with the
cupula of the sucker and the tentacle with the caruncle. This interpretation was supported
by Professor Lankester (1883, Encijc. Brit), and is in feet widely held'. For my part I am
unable to subscribe to the very ingenious theory of Valenciennes for reasons which I shall
submit later. Valenciennes not only compared the sheathed tentacles of Nautilus with
the acetabula of Dibranchs, but grouped them together in such a way as to show
a correspondence with the eight arms of an Octopod. His point of view was undoubtedly
ingenious though artificial. In his PI. xi. fig. 2, he represents an ideal section across
the cephalopodium, in which the tentacles are associated together into four paired groups
represented, as he claims, in actual anatomy by eight muscular peduncles. These groups
are as follows : —
1. Hood and accessory sheath.
2. Mass of the external digital lobe.
3. Internal labial lobe.
4. Inferior labial lobe of $ .
Valenciennes explained his position in these words (pp. 27.5 — 276): — "Dans cette
maniere de voir je ramene au nombre ordinaire des bras des c^phalopodes en general
le nombre de ceux de nautile, puisque les quatre-vingt-huit cirrhes, consideres avant moi
comme bras du nautile, ne sont plus que des appendices sortant des ventouses allong^es
en gaines Mais ne voulant pas pousser ces analogies au-dela de ce qui peut
en quelque sorte se demontrer, je me hate d'ajouter que si les bras du nautile ressemblent
a ceux du poulpe par leur position autour du bee, ils sont toutefois bien diff'^rents par
leur forme et leur structure, malgre I'espece d'analogie que nous venons d'indiquer entre
les cirrhes de I'un et les ventouses de I'autre."
Not the least noteworthy feature in regard to the digital tentacles of Nautilus,
especially those of the outer whorl, is their striking constancy in number and position.
This is the more remarkable because their number is large and their position apparently
arbitrary. Owing to the latter circumstance it has not been easy to arrive at a satis-
factory method of enumeration of the tentacles, which is a matter of statistical interest
and incidentally of morphological importance.
The sheaths of the tentacles cohere in such a manner that they form a fieshy mass
in which the tentacles themselves appear to be arranged without regard to any law or
order, but a critical examination discloses, to my thinking, a possible means of inter-
pretation which the diagrams on Plate LXXXI. will serve to elucidate.
The method of enumeration which I have to suggest was acquired independently
after many attempts to amve at a settlement. Subsequently I became acquainted witii
' /','.(/. it is the view expressed in Professor Arnold Lang's Lehrbiich der veryUichenden Aiuitomie, Ist edit.
1892, p. G91.
770
DIGITAL AND OPHTHALMIC TENTACLES.
Professor Vayssifere's efforts in this direction'. As it is, in my opinion, a subject of
considerable relative importance, I will venture to quote the passage from M. Vayssiere's
memoir bearing upon this point : — " S'il est facile de numeroter les bras d'un Cephalopoda
Dibranchial vu leur nombre restreint, il n'en est pas de meme pour ceux du Nautile ;
il est incontestable que je designerai sous le numero 1 celui qui forme la moitie du
capuchon cephalique, c'est lui qui est le plus dorsal. Le numero 2 pourra s'appliquer
au premier tentacule qui longe le bord du capuchon en contractant adherence avec lui
sur presque toute sa longueur; mais pour les autres quel ordre faudra-t-il suivre pour les
numeroter ? est-ce un peu au hasard, en allant de dedans en dehors et en s'avan^ant
progressivement vers I'entonnoir, ou bien faut-il tenir compte de certains groupements de
trois a quatre tentacules qui forment des sortes de verticilles se recouvrant ? Ce dernier
mode serait prefei-able si Ton ne constatait pas quelques variations dans la disposition
de ces groupes ; aussi vaut-il mieux s'en tenir a la numerotation de la face dorsale vers
la face ventrale, en allant toujours par rangees trans versales de I'int^rieur vers I'exterieur."
The desirability of a definite method of enumeration by which the indi\'idual members
of a relatively constant series can be recognised, will be obvious to all who interest
themselves in meristic phenomena, and especially to those who may have opportunities
for studying Nautilus.
The following tabulations display the differences in the application of my method as
compared with that of Professor Vayssiere, and I hope to show by what has preceded as
well as by what shall follow, that this is a point about which it is worth while to differ.
Table A.
my standard.
Enumeration of the tentacles of the outer circlet in Xautilus according to
Wn.LEY
1 .
2
3 .
4 .
5 .
6 .
7 .
8 .
9 .
10 .
Vayssi£be^
1
2
3
.. 5
7
.. 10
.. U
.. 19
.. 16
.. 12
WlLLEY
11 .
12 .
13 .
14 .
15 .
16 .
17 .
18 .
19 .
VayssiSre-
. 9
. 6
. 4
8
. 11
. 13
. 18
. 15
. 17
Table B. Enumeration of the outer tentacles according to Vayssiere's standard.
VAYSSlfeBE
1 ..,
O
3
4
5
6
7
8
9
10
WiLLEY
1
2
3
13
4
12
5
14
11
6
VAYSSlfeRE
11
12
13
14
15
16
17
18
19
WiLLEY
. 15
. 10
. 16
7
. 18
. 9
. 19
. 17
. 8
1 Vayssiere, A., " Etude sur I'organisation du XautUe." Ann. Set. Nat. (8) n. 1896, see p. 162 and PI. 17, fig. 8.
2 Vayssiere, op. cit. PI. 17, fig. 8. The figure relates to a male N. macromphalus.
DIC4ITAL AND OPHTHALMIC TENTACLES. 771
The above tables seem to betray ineradicable divergencies, due merely to different
points of view. There is, however, one door left open for immediate criticism. It might
at least have been expected that there would be agi-eement as to the first and the
last, but such is not the case. Vayssiere's nineteenth tentacle is my eighth, and my last
is Vayssiere's seventeenth.
It was admitted by Owen and his successors that the main bulk of the hood consists
of "two digitations conjoined along the mesial line'." "The median antero-posterior line
traversing this hood exactly corresponds to the line of concrescence of the two halves
of the fore-foot, which primitively grew forward, one on each side of the head, and finally
fused together along this line'." Seeing that the lateral portions of the hood are actually
formed by the cohesion of the sheaths of the second pair of tentacles with the main
body of the hood, it appears reasonable to infer that the latter arose by concrescence
of the sheaths of the first pair. The median portion of the hood of Nautilus is thus
taken to represent a dorsal symphysial tract.
I have found no reference to the occurrence of a ventral symphysial tract'^ and yet,
I think, that is the nature of the bridge which forms the fundus of the concavity which
lodges the funnel, and at the same time completes the outer circlet (annular lobe of
Lankester) across the ventral median line. At the corners of this tract are placed the
tentacles of the nineteenth pair, in very much the same way as the tentacles of the
first pair occur near the outer borders of the median portion of the hood, and I think
it likelv that this ventral tract owes its origin to the concrescence of the sheaths of the
last pair of tentacles, just as the dorsal tract admittedly represents the fused sheaths of
the first pair. An indication of the bilateral constitution of the ventral tract is furnished
by the mesial line which divides the laminae of the mucous gland which develops upon
its inner surface in the female*. If it is true that there is a ventral symphysial tract, then
it is clear that the tentacles which are most intimately associated with it and immediately
border it are to be reckoned as the last of the series.
Assuming then that my identification of the nineteenth pair of tentacles of the outer
circlet is correct, it still remains to justify the enumeration of the tentacles which
intervene between the first and the last. Taking into consideration their evident character
as marginal appendages of an annular lobe, we may further assume that the border out
of which they arose was not plain but fi-illed like an undulating membrane, and that
1 Owen, Memoir 1832, p. 13. Owen seems to have erred somewhat iu his counting of the tentacles
through anxiety on behalf of even numbers. The passage runs as follows: — "Of the digital processus nineteen
have been enumerated on either side ; but as the hood has two perforations anteriorly from wliich tentacles
similar to those of the digitations are also protruded, we may consider this part as two digitations conjoined
along the mesial line, and so reckou with Ruraphius the even number of twenty digitations on either side
of the head."
- Lankester, E. R., Article " MoUusca," Encycl. Brit., 9th edit., republished in Zoological .4rticli's, 1891, see
p. 137.
^ Except perhaps in the memoir by Valenciennes (1841), who describes the two inferior arms as being
united below by a thin plate hollowed out to receive the funnel, while on the inner surface of the plate,
on each side of the median line of junction, there is in the female a lamellated organ, the same which
has recently been shown by Mr Graham Kerr to serve as a receptacle for the spermatophore.
■* Cf. Kerr, J. G., " On some points in the Anatomy of Nautilus pompilius." Proc. Zool. Soc. I.ond.
1895, see PI. 39, fig. 3. I shall subsequently refer to this laminated gland as the " Organ of Valenciennes."
w. VI. 101
772 DIGITAL AND OPHTHALMIC TENTACLES.
the sinuations underwent concrescence, thus producing a thick matrix in the midst of
which the tentacles occur, in places, three or even four deep. By slicing off the upper
portion of the sheaths of the outer tentacles, it may be at once seen that the sequence
of the numbers one to eight and nine to thirteen, in my enumeration, is exactly such as
would result from the concrescence of a frilled border, and closer examination will show
that the remaining numbere follow suit in the order given.
Not only are number and plan of distribution of the outer tentacles identical
in male and female, but they are also the same in the three species, iV. pompilius,
N. inacromphalus, and 3'. umhilicatus. In all cases the outermost of the series are numbers
5, 6, 7, and 8, of which the first three appear to be grafted upon the external surface
of the circlet, and are much shorter than the rest of the tentacles whose sheaths form
the outer wall of the crown, number 6 especially lagging far behind the others, its sheath
being produced into a prominent lobe behind. The prism-shaped sheath of the 7th tentacle
is always accurately wedged into a deep gi-oove between the 9th and 10th ; that of the
6th fits into a similar groove between the 5th and 11th, while the 5th itself is ensconced
at a higher level between the 4th, the 12th and the 11th. There is a shallow concaWty
between the sheaths of the 9th and 17th tentacles which receives that of the 8th;
the inner border of this concavity is furnished by the sheath of the I7th, and so
shuts off the 9th from participation in the lateral wall of the great ventral conca\'ity
of the annular lobe which I have called the fossa infundibularis ; this wall is actually
formed by the coalescent sheaths of the 8th, 17th and 19th tentacles.
If we now look for some corroboration of the sequence of tentacles which I have
suggested above, from a consideration of the nerves which are distributed among them,
we shall find to a certain extent a welcome confirmation. My series 5 — 8 inclusive,
represented in Professor Vayssiere's scheme by the numbers 7, 10, 14 and 19, are found
to receive their nerves in absolute sequence (PI. LXXXI. fig. 6, and PI. LXXXII. fig. 6).
The most direct manner of procedure for exposing these nerves, and I believe the
technically correct method, is to remove one of the alae infundibali, and then by
horizontal incisions through the cartilage and coriaceous tissue at the base of the
cephalopodium gradually to expose the sense-centres of one side. In this way it is
impossible to overlook the otocyst which escaped both Owen and Valenciennes, being
first found by Macdonald (1855, Phil. Trans.). In front of and partially concealed by
the otic capsule appears the dark mass of the pedal ganglion. Upon exposing this portion
of the "sjTicerebrum'" without too much displacement of neighbouring organs, three
nerve-roots almost simultaneously appear to the view of the dissector, and upon tracing
these to their destination I was somewhat agreeably surprised to find that they were
the nerves belonging to the sixth, seventh and eighth digital tentacles according to my
system of enumeration (PL LXXXI. fig. 6). Continuing the dissection by turning aside
the olfactory labyrinth (rhinophore), we find that contiguous and level ^-ith the root of vi
is that of V (PL LXXXII. fig. G). The large nerve lying at a slightly deeper level
next to V in the dorsad direction is iv, at the base of which the root of ill arises
apparently as a branch, but the apj^earance of branching is attributable to the fact that
* The term applied bj Professor LaDkester to a compound cephalic ganglionic mass iu contrast with
" archicerebrum " where the cephalic centre comprises the cerebral ganglia only.
DIGITAL AND OPHTHALMIC TENTACLES. 773
the two nerves are for a very short distance enclosed in a common perineurium. The
next nerve dorsad of in and IV is ii, which has a length of about 21 mm. from its origin
to its insertion into the base of the second tentacle. Then comes i with a length of
28 mm. giving off collateral branches to the substance of the hood. By cutting through
and turning back the roots V, vi, vii and viii, those of IX, x, xi and Xii are exposed
in sequence at a deeper level, ix occurring between vill and Vii, x between vii and
VI, XI in front of vi ; xii lies at a slightly deeper level, and bears the same relation
to XI that IV does to v. The sequence of IX, X and XI is as striking as that of
V, VI, VII and viii but I am inclined to lay special stress upon the latter sequence.
The root of xvii lies opposite to viii but at a deeper level than the series IX, X and XI.
Without attempting the impossible task of proving to complete satisfiiction the
truth of the entire sequence suggested by me, I think it is legitimate to select a small
group as a test case, and in this sense I may repeat the fact, which is not at all
obvious without actual demonstration, that the nerves v, vi, Vii and viii occur in
indubitable topographic sequence (PI. LXXXII. fig. 6).
Before devoting a few lines to a consideration of the inner whorl of the cephalo-
podium, it is necessary to turn our attention to the ophthalmic tentacles which occur in
front of and behind the eye. These resemble the digital tentacles in shape and annulation,
but differ, in that the sulci are much deeper on the side corresponding to that on which
the suctorial ridges of the former are placed, so that the tentacles are converted into
distinctly lamellate appendages, and when examined in the fi'esh state under the micro-scope,
the grooves and lamellae are found to be covered with vibratile cilia (PL LXXIX. fig. 14).
The suctorial ridges of the outermost digital tentacles may in favourable specimens
be seen to become more and more prominent and lamelliform towards the free ends,
and I have observed that when the main portion of the tentacles has been affixed
to a surface, the tips of the tentacles which probably exercise some kind of tactile
function were free. I have never seen cilia at any part of the true digital tentacles.
The ciliated lamellae of both the pre-ocular and the post-ocular tentacles are directed
forwards, and the shafts of the tentacles when extruded are kept in an attitude of
attention, the anterior being directed obliquely outwards and forwards, the posterior
obliquely outwards and backwards, but nearly or quite straight (cf PI. LXXVII. figg. 8, 6).
I came to the conclusion' that the ophthalmic tentacles of Nautilus while homologous
with the adhesive digital tentacles have become modified to serve as accessory olfactory
organs, and that the ciliated ridges represent olfactory lamellae. In the living state the
ophthalmic tentacles afford a strong contrast to the brown digital tentacles by reason of
their nearly uniform white colour ; there is a small trace of brown pigment in the
annulations and at the edges of the lamellae, but when viewed under water the general
colour is white. They are also characterised by increased sensibility. Whereas the digital
tentacles can be touched without being retracted in consequence, the slightest contact
with a foreign body occasions the instant withdrawal of the ophthalmic tentacles into
their sheaths.
1 Willey, A., "The pre-ocular and post-ocular tentacles o{ Nautilus." QiDirt. J. ilicr. Sc. Vol. 40,
1897, p. 199.
101—2
774 DIGITAL AND OPHTHALMIC TENTACLES.
In my paper on this subject (1897) I added the following remarks: — "The presence
of accessory olfactory tentacles in Nautilus can, I think, be related to an essential
bionomical difference between the existing Tetnibranchiata and the Dibranchiata. Nautilus
finds its food chiefly by the sense of smell, while the Dibranchiata with their
remarkably perfect eyes pursue their quarry by the sense of sight. This difference
is further emphasised by the different modes adopted by the natives for trapping these
animals.
" One of the surest ways of obtaining Nautilus, and in fact the method by which
I have obtained most of my specimens at Lifu, is to bait the fish-basket with the
cooked and bruised exoskeleton of Palinurus wrapped up in cocoa-nut fibre like
a small parcel and placed in the fish-trap overnight. There is nothing to be seen but
something to be smelt, and by this means I have obtained as many as ten
Nautilus at one time. For taking Octopus the natives of Lifu employ a very different
method. A rounded oval piece of stone backed by a well-fitting piece of the shell of
a species of Cypraea, to which are added pieces of leaf to simulate legs and tail, is
dangled along the surface of the water at the end of a line. The natives say that
the Octopus mistakes this for a rat, agamst which it has a special grudge; but
the fact remains that Octopus attacks this singular non-scented contrivance and so is
captured." The Lifuan name for this simple apparatus is " athy."
One of the most interesting points about the ophthalmic tentacles concerns theii-
innervation, regarding which there has been serious disagreement among zoologists, but
I hope to show in this matter that the facts of the case are beyond the reach of
controversy. Owen's figure (1832) of the central and peripheral nervous system of
Nautilus conveys an excellent general idea of its main characters, but is too diagram-
matic in its representation of details to be of much assistance at this particular juncture.
Valenciennes (1841, p. 288, PI. viii. fig. 2) described the nerves of the ophthalmic
tentacles as arising from the cephalic ganglionic ring, separated fi-om each other on each
side by a wide interval which is occupied by the optic and true olfactory nerves, so
that according to him the nerve which supplies the posterior ophthalmic tentacle arises
dorsad and mediad of the optic nerve. Valenciennes appears to have overlooked the
buccal or stomatogastric nerves, of which there are two pairs arising fi-om the cerebral
(supra-oesophageal) commissure, and hence probably misinteqireted one of the latter as
the nerve to the posterior ophthalmic tentacle (cf PI. LXXXIII. figg. 28 and 29).
Nevertheless von Jhering", who was the first to note the existence of a stomatogastric
or " sympathetic " nervous system in Nautilus, confirmed Valenciennes' account of the nerves
to the ophthalmic tentacles, which we may be allowed to call the ophthalmic nerves in
place of a better name although it is of course open to misinterpretation ; but I am
not prepared to suggest a new name. Von Jhering says that the posterior ophthalmic
nerve is the last nerve which leaves the supra-oesophageal ganglion dorsad of the optic
nerve. As a matter of fact this last cerebral nerve is the second buccal or stomato-
gastric nerve which von Jhering overlooked. With regard to the anterior ophthalmic
' Jhering, H. v., " Vergleichende Anatomie des Nervensystemes und Phylogenie der Molhisken," Leipzig,
1877, pp. 262 and 264. Von Jhering's account of the stomatogastric system has been essentially corrected and
amplified by Mr J. Graham Kerr {op. cit. 1895), whose description of these nerves I can confirm.
DIGITAL AND OPHTHALMIC TENTACLES. 775
nerve, von Jhering also confirmed Valenciennes in these words : — " Nach innen und unten
vom Sehnerven entspringen vom oberen Theil des vorderen Suboesophagealstranges der
Geruchsnerv und der Nerv des vorderen, unteren oder ventralen Augentakel." This
of course is a very broad statement, and in fact the true olfactory nerve does lie below
the optic ganglion, but it does not arise from the anterior (pedal) commissure (cf my
PI. LXXXI. fig. C).
Pelseneer' regarded the ophthalmic tentacles of Nautilus as belonging essentially to
the head, and as homologues of the labial and nuchal cephalic tentacles of Gastropoda,
and therefore not honiodynamous with the digital tentacles of Nautilus. While advocating
the theory of the pedal nature of Cephalopod arms in general, he insists on the cephalic
nature of the ophthalmic tentacles of Nautilus, saying in a foot-note : " II est inexact de
dire, comme von Jhering, que le tentacule ophthalmique anterieur est innerve par le
ganglion infra-oesophagien anterieur. Tout comme le nerf olfactif, le nerf du tentacule
ophthalmique anterieur sort de la partie laterale extreme du ganglion supra-cesophagien,
ainsi que le montrent les figures precit^es de Valenciennes."
It is therefore quite clear that the ophthalmic tentacles of Nautilus occupy a crucial
position as between head and foot, and it is the more desirable to ascertain the exact
origin of their nerves. The most recent author who has dealt with this matter is
Dr L. E. Griffin'^, who describes an accessoiy nerve-trunk associated with the axial nerve-
trunk traversing the body of the tentacle, and adds that " the nerves of the two ocular
tentacles of each side are branches of a nerve which comes off from the pedal ganglion
near the outer end, which also sends branches to the hood."
According to my own observations Dr Griffin's statement comes nearer to the truth
than any other relating to this point with which I am acquainted, but I do not agree
with him that the ophthalmic nerves are branches of the hood-nerve.
The ophthalmic nerves in fact arise by independent roots from the dorsal boi-der
of the pedal ganglion'. The anterior ophthalmic nerve arises close behind the capitular
(hood) nerve, but is surrounded by its o\vn perineurium, and proceeds upwards and
slightly forwards to the base of the tentacles (PI. LXXXII. fig. 6). The posterior
ophthalmic nerve again arises independently, close behind its colleague and holds
a pronounced recurrent course, passing backwards to its tentacle in the wall of a sinus
in which the optic ganglion lies. The posterior ophthalmic nerve does therefore not arise
dorsad of the optic nerve, but it passes backwards from its origin in the pedal ganglion
(PI. LXXXII. fig. 6). The position of the ophthalmic nerves in relation to the digital
nerves shows that the ophthalmic tentacles are in sequence with the digital tentacles,
and that the posterior ophthalmic tentacle is the first, the anterior ophthalmic tentacle
the second, and the capitular (first digital) tentacle the third member of a continuous
morphological series, the remaining digital tentacles following in their due order.
In the diagrams of transverse sections on PI. LXXXI. the ophthalmic tentacles are
' Pelseneer, P., " Sur la valeur morphologique des bras et la compositiou du systfeine nerveux central des
C^phalopodes." Arch. Biol. Vol. viii. 1888, see p. 730.
- (irillin, L. E., "Notes on the tentacles of Nautilus p(impiiiu,v." Johns Hopkins Univ. Circ. Vol. xviii.
Nov. 1898.
' For the present I use the term "pedal" in this connection as being less unwieldy and less inaccurate
than " infra-oesophageal." It might be called by an indifferent name, such as " coronal ganglion." " Brachial
ganglion" implies too much. "Tentacular ganglion" would also mislead.
776 DIGITAL AND OPHTHALMIC TENTACLES.
omitted, and I have also not reckoned them numerically with the digital tentacles, but
their morphological valency as the fii'st two tentacles of the outer whorl of the cephalo-
podium of Nautilus is I think clear, and I shall again refer to the recurrent nerve
of the posterior ophthalmic tentacle'.
The inner whorl of the cephalopodium is incomplete dorsally, and moreover it is
broken up into distinct tentaculiferous and lamelligei'ous lobes which have become more
or less specialised as accessory sexual organs, and exhibit profound sexual differences
which are tolerably familiar to malacologists. There are two principal groups of tentacles
of the inner whorl which may be described as extrahuccal and infrabuccal, and correspond
to Owen's " external labials " and " internal labials " respectiveh'. The objections to Owen's
terminology are firetly that the epithets " external " and " internal " obscure the view that
the tentacles belong to an inner whorl, and secondly the expression " labial " is inappropriate
and apt to cause confusion with the labial papillose ft'inge which surrounds the jaws.
In the female the exti-abuccal lobes carry twelve ordinary tentacles on each side,
of which eight are disposed in a descending and four in an ascending series. The lobes
themselves consist of the coalescent sheaths of the tentacles borne upon a fleshy basis
which rises up from the fundus of the fossa buccalis. The extrabuccal lobes adhere
ventrally to the inner surface of the outer whorl- by means of an intercoronal membrane
which proceeds on each side from the ventral border of the lobe (cf PI. LXXX. figs. 7
and 8, i).
In the male the extrabuccal tentacles are subdivided into three groups, an upper
series of eight tentacles with coalescent sheaths, a middle set of three, and a single
ventral fi-ee tentacle. The free tentacle is usually found to nestle within a groove formed
by a vertical flap which grows out from the sheath of the first tentacle of the middle
set or the ninth of the entire series, or it may follow simply at the end of the series.
The middle set comprising the ninth to the eleventh extrabuccal tentacles, together with
the free twelfth tentacle, constitute a natural group of four which are known as the
spadix on the one side and the antispadix on the other. The intercoronal membrane
arises in the male from the lower border of the upper set of coalescent tentacular sheaths
(1 — 8), and passes unattached in front of the tetrad group to its insertion into the
inner wall of the outer (annidar) lobe.
The ventral or infrabuccal lobe of the inner whorl is modified in both sexes to form
a special infrabuccal apparatus. In the female it consists of a median laminated organ
flanked on each side by a uniserial row of 12 — 14 tentacles; the lamellae are modified
tentacles and retain their apices (PI. LXXIX. fig. 6, and PI. LXXX. fig. 11). In the
male there are no unmodified tentacles remaining in connection with the infrabuccal
apparatus, which is completely metamorphosed into a laminated organ, known as Van der
Hoeven's organ, retractile within a pouch and closely adherent to the ventral integument
of the buccal cone.
I do not propose to enter upon a long description of these structures, partly because
this has already been done by others, and partly because I was unable to ascertain the
manner in which the various parts are brought into action.
' In my experience the most suitable material for the successful dissection of the tentacular nerves of
Nautilus consists of soft specimens and not of such as have become too hard under preservation.
- The outer wborl is the "oral sheath" of Owen, the "annular lobe" of Lankester.
DIGITAL AND OPHTHALMIC TENTACLES. 777
The spadix is a composite erectile organ (PI. LXXVIII. fig. 4) which occurs indiiferently
on the right or left side of the cephalopodium. It was originally described by Van der
Hoeven', and has since been treated in greater detail by Professors Haswell* and Vayssiere'.
This organ shows a gradual development from the early to the adult life of the animal,
only attaining its full growth at se.xual maturity (cf PL LXXX. figg. 3 — 7 and PI. LXXIX.
figg. 9 and 10). The ninth extrabuccal tentacle constitutes the glans and its sheath, the
prepuce. The latter gives rise to a free fold along its outer border, and at the summit
bears a large thick glandular cushion covered with bordered pits, which may be described
as the preputial (/land. The tenth tentacle (the second of the spadix) is a fleshy flattened
appendage strongly ribbed upon its glandular surface. The third tentacle of the spadix
(eleventh of the series) is also a flattened foliaceous appendage, but in place of the
transverse costae of the preceding tentacle, it presents a marvellous system of glandular
crypts, producing the appearance, as Professor Haswell pointed out, of a minute honeycomb.
The last tentacle, the twelfth extrabuccal, is somewhat enlarged, but otherwise undergoes
comparatively little modification and is partially concealed, on the outer aspect of the
spadix, beneath the flap of the prepuce. The second tentacle of the spadix may be
distinguished as the costate tentacle, the third as the tessellate tentacle, and the fourth as
the free tentacle or cirrus.
Like all the other accessory sexual organs of both male and female, the spadix manifests
a gradual development pari passu with the gi-owth of the animal. I use the word
" development " in contrast with " growth " to express the fact that in the young Nautilus
the character of the spadix is quite different from the adult structure of the organ.
The smallest Nautilus which I ever obtained was a male iV^. pompilius* with the
followng dimensions : —
Length from root of siphuncle to mid-anterior point of hood (measured
along the dorsum)' 25 mm.
Length of hood along middle lino 10'5 „
Breadth of body across middle of eyes ...; 15 „
The shell was perforated at the umbilicus and contained thirteen air-chambere.
At this stage it is not possible to say upon which side of the cephalopodium the
definitive spadix will appear, the two groups of four tentacles — numbers 9 to 12 of the
inner whorl — being almost exactly alike, notably in regard to the tardy development
of the 10th tentacle (PI. LXXX. figg. 1 and 2).
After this stage the growth in length of tentacles 9 — 11 of the future spadix becomes
' Van der Hoeven, J., "Bijdragen tot de ontleedkundige kennis aangaande Nautilus pompilius L." Verh.
koii. Akad., Amaterdanj, 1856.
' Haswell, W. A., " Note on certain points in the arrangement and structure of the tentaculiferous lobes
in Nimtilm pompilius." P. Linn. Soc. N. S. Wales, (2) x. 1895, p. 544, PI. 48.
' Vayssi^re, A., up. cil., Ann. Sci. Nat. (8) ii. IHSIG.
■• Willey, A., " In the Home of the Nautilus." Natural Science, Vol. vi., June 189.5, p. 410. The surface
of the hood was white and unpigmented when the specimen was brought to me, but this may have been
due to previous discharge of pigment from the epidermal cells of the hood. This often happens when the
animals are kept uuder unfavourable conditions.
° This length in an average adult male attains to about 140 mm. I have seen much larger shells of
N. pompilius from the Moluccan Sea than any that I obtained in New Britain.
778 DIGITAL AND OPHTHALMIC TENTACLES.
arrested while number 12 continues to grow, and thus appears for a time to be the
most important member of the confederation, until eventually its limit is reached, the
others forge ahead and number 12 retires behind the curtain which grows out from the
sheath of number 9 (cf PL LXXX. figg. 7 and 4)'.
The infrabuccal organ of the male consists of a bipartite laminated structure contained
in a pouch formed by the integument of the buccal membrane, which in adult specimens
projects far back into the peristomial haeniocoel (PI. LXXXII. fig. 3). It was first
described and figured by Van der Hoeven- (1850) who regarded the lamellae as tentacles
modified in a manner similar to those of Owen's laminated organ in the centre of the
infi'abuccal lobe of the female.
Van der Hoeven's organ (PL LXXX. figg. 13 — 15) has recently been described in
some detail by Dr L. E. Griffin^ It consists in the adult of a pair of fleshy narrow
lobes parallel to one another, closely approximated in the middle line and bearing, upon
the outer surface of each about fifteen to nineteen lamellae of which only a certain
number are visible in external view, the others being concealed below them like the
leaves of a book.
The lamelligerous lobes are united together at the base by an intermediate portion
which forms a sort of peduncle, and the adoral surface of lobes and peduncle is beset
with the openings of glands, which discharge their secretion into the cavity of the pouch
in which the organ is contained (PL LXXX. fig. 14). Often in preserved specimens
a coagulum is found at the orifice of the pouch. In sections cutting the lamellae at
right angles the epithelium clothing the surfaces of the lamellae appears to be richly
ciliated, an appearance due, as described by Griffin, to the occurrence of numerous
sensory cells bearing sense-hairs.
In a very young male of X. pompilius, the smallest specimen which I obtained, the
appearance of the infrabuccal organ is different from that which it presents in the adult.
Instead of being bent into a U -shape as it becomes at a later stage, the two halves of
the gland are extended in the transverse direction with a narrow median tract between
the ridges which represent the future lamellae (PL LXXIX. fig. 12). The appearance of
the infrabuccal organ at this stage suggests that its lamellae are homologous with the
tentacles only of the female infrabuccal organ, while the group of lamellae (Owen's
"olfactory" organ) between the tentaculiferous lobes is reduced to an insignificant median
tract in the male. If this be so, then the infrabuccal organ of the male is not simply
homologous with that of the female, but chiefly with the tentacular portion of the latter,
the median lamellae of the female organ being mostly suppressed in the male, and the
lateral tentacles of the former converted into lamellae in the latter. Apart, however, from
the interesting evidence furnished by the development of the male infrabuccal organ, its
composition in the adult suggests the same homologies which we may tabulate.
' The spadix of a young Nautihix of an age corresponding to that represented on PI. LXXX. figg. 4 — 6,
has been described and figured by Prof. Vayssiere op. cit. 1895, PI. 16, figg. 3 and 4, and PI. 18, tig. 14.
= Trans. Znol. Snc. London, Vol. rv. Part 1, 1850, PI. 8, fig. 9.
' Johns Hopkin.'i Univ. Circ. xviii. November 1898, p. 12.
DIGITAL AND OPHTHALMIC TENTACLES. 779
Homologies of In/rabuccal Organ of Male and Female Nautilus.
Male. Female.
Lamelligerous lobes Tentaculiferous lobes
Peduncle Lamelligerous lobe
I cannot say with certainty whether the innervation of the infrabuccal apparatus
of male and female affords conclusive evidence of the truth of the above comparisons,
but I think it may.
The first accurate account of the innervation of the female infrabuccal organ was
furnished by Mr Graham Ken- in 189.5'. A single nerve-trunk, the last to leave the
pedal ganglion, external to the infundibular nerve (PL LXXXII. figg. 4 and 5) crosses
the peristomial haemocoel on each side, and penetrates into the base of the infi-abuccal
organ. " This bends towards the middle line, pursues a curved couree in the substance of
the lobe, and meets with its fellow of the opposite side " (Kerr, op. cit, p. 676). From
this commissural nerve, which I propose to call after its discoverer, Kerr's collector
or commissure, branches are given off which pass directly to the tentacles and lamellae
individually. I have dissected out this remarkable nerve in the female infrabuccal organ,
but in the male I have been unable to trace a connection between the right and left
infrabuccal nerves, the median portion of the nerve from which, in the female, the branches
to the lamellae arise, apparently not occurring in the male (PI. LXXXII. figg. 1 and 2).
Finally it may be remarked that the infrabuccal apparatus of Nautilus in general
combines the properties of gland and sense-organ, but that it is in any case an accessory
sexual organ since it differs in male and female. It is therefore not appropriate to
ascribe an olfactory function to it, since this e.xpression is too precise in the present
state of our knowledge and moreover leads to confusion with the true rhinophore^ and
the ophthalmic tentacles which present no sexual differences.
There is another lamellar organ in the female to which I have already referred,
which occupies the inner surface of what I have called the ventral symphysial tract.
It is composed of two halves separated in the middle line, and the lamellae of each
half are disposed in an arcuate manner, running at first longitudinally, and then curving
inwards or transversely to the middle line.
This organ was first correctly described by Valenciennes^, and I shall call it the
Organ of Valenciennes, in order to distinguish it from the infrabuccal lamellar organ of
the female which is the Organ of Owen, and from the infrabuccal lamellar organ of
the male which is the Organ of Van der Hoeven.
The median division which separa,tes the two halves of the organ of Valenciennes
is not always very distinct, and Kerr'' has figured an example in which the lamellae
' Kerr, J. G., op. cit., P. Zool. Soc. London, 1895, p. 675.
'' Owen (1832) surmised that the lamellae of the female infrabuccal organ exercised an olfactory function,
in ignorance of the sexual differences which were first made known by Van der Hoeven (1850) and in
ignorance of the rhinophore which was discovered by Valenciennes (1841).
^ Valenciennes, A., op. cit. 1841, p. 277, PI. xi. fig. 1 (.
* Kerr, J. G., op. cit. 1895, P. Zool. Soc. PI. xxxix. I have also observed that some of the lamellae
may become confluent across the middle line, while others in the same organ remain distinct on each side.
w. VI. 102
780 PERISTOMIAL HAEMOCOEL ; SYSTEMIC AORTA ; CEPHALIC ARTERIES.
were continued uninterruptedly from one side to the other. KeiT has further found
a spermatophore adherent to the lamellae of this organ, thus indicating its function as
a receptacnlum seminis. Curiously enough I have not found a spermatophore in this
position in any of my preserved females, but in a living female I once observed the
end of a spermatophore protruding from the circumoral complex as shown in fig. 2,
PI. LXXYII. I placed this specimen carefully back into its cage, in the hope of obtaining
some result, but nothing came of it.
On Plate LXXX. figs. 9 — 12, I have rendered all the secondary sexual characters
of a young female, measuring 66 mm. in length from the root of the siphuncle to the
anterior border of the hood. The organ of Valenciennes is mapped out but barely laid
down, although foint traces of the future lamellae are discernible at the sides. The
extrabuccal lobes (PI. LXXX. fig. 10) exhibit normal development, since they are not
specially modified in the female, whereas the infrabuccal lobe shows a retarded develop-
ment, small in size and with the tentacles just appearing (fig. 11). In conclusion it
may be noted, although not coming within the scope of the present chapter, that the
nidamental gland is only represented by an inconspicuous primordial tract on the inner
surface of the ventral mantle-flap (PL LXXX. fig. 12).
10. Peristomial Haemocoel; Systemic Aorta; Cephalic Arteries.
The venous system of Nautilus consists of definite venous channels, of which the
most conspicuous are the vena cava, the pallial veins and the siphuncular vein, of inter-
stitial lacunae and of spacious sinuses. The sinuses are traversed by muscles, nerves,
and conjunctive trabeculae, and form vast cisternae, surrounding the buccal cone, the
crop, and the liver. These three principal dinsions of the haemocoel may be distinguished
as the peristomial, peri-oesophageal and peri-hepatic haemocoels respectively. They com-
municate by separate apertures -n-ith the vena cava, which collects all the blood and
transmits it through the central sinus venosus and afferent branchial vessels to the gills.
The peri-oesophageal haemocoel communicates with the vena cava by means of the
numerous fenestrations in the dorsal wall of the latter which were described and figured
by Owen (cf my PL LXXXI. fig. .5).
In preserved specimens the liver is often surrounded by an abundant coagulum which
conceals the hepatic lobules from view. The wall which shuts off the peri-hepatic sinus
from the perivisceral coelom is a thin transparent membrane. The liver, as is well known,
consists of right and left portions, and the peri-hepatic haemocoel accordingly opens into
the sinus venosus by two orifices which lie mediad of the posterior afferent branchial
trunk (which supplies the smaller gill on each side), and dorsad of the genital duct
on the right side, and of the pyriform body on the left. These orifices are seen at
the back of the sinus venosus after cutting across the rectum which traverses the latter.
The peri-hepatic haemocoel is nevertheless nothing more than a backward prolongation
of the peri-oesophageal coelom, protruding like a hernia into the perivisceral coelom. That
this is the case may be seen at once by cutting into the haemocoel from the dorsal
PERISTOMIAL HAEMOCOEL ; SYSTEMIC AORTA; CEPHALIC ARTERIES. 781
side, when a forest of slender conjunctive trabeculae are found passing from the inner
(mesial) surfiice of the liver to the wall of the crop.
The peristomial haemocoel is virtually separated from the peii-oesophageal space by
a tendinous diaphragm on the dorsal side, and by the tissues and organs of the fundus
elsewhere. The diaphragm is a very definite structure stretching from the cerebral
capsule to the body-wall in the region of the nuchal membrane (PL LXXXI. fig. 7, and
PI. LXXXIII. fig. 26).
The nerve-collar which strictly surrounds the oesophagus is the cerebro-visceral loop.
The pedal commissure is separated from contiguity with the oesophagus by an interval
which is occupied by the ventral retractor muscles of the buccal mass. The oesophagus
passes into the buccal mass precisely at the level of the fundus of the peristomial
haemocoel, so that the oesophagus proper lies entirely within the peri-oesophageal haemocoel,
the buccal mass' with its attendant muscles and nerves being the only portion of the
digestive apparatus which occurs within the limits of the peristomial haemocoel.
The cerebral jjortion of the cerebrc^-visceral nerve-collar lies within the peristomial
haemocoel (PI. LXXXI. tig. 7) while the vis-
ceral portion lies within the peri-oesophageal
haemocoel (PI. LXXXIII. fig. 26). Within
the mass of the cephalopodium at the angle
of junction of the cerebral and visceral por-
tions of the nerve-collar occur the cephalic
organs of special sense (eye, rhinophore, and
otocyst) and their nerves. The demarcation
of these components of the nervous system is
therefore clear.
The greater or systemic aorta of Nautilus
arises as a large truncus arteriosus'^ from the
left dorso-posterior region of the heart and
leaves the pericardium through the left peri-
cardio-visceral fontanelle. On its mesial side
as it leaves the heart it is adherent to the
ligament which binds the heart to the floor
of the pericardium. Haller (1895) points out
that this cardiac ligament is a very primitive
structure, representing the place of origin of
the heart in the coelom.
Having emerged from the pericardium,
the great aorta, after a short upwardly directed
course through the perivisceral coelom, enters the peri-hepatic haemocoel between the
■ There are no extra-bulbar salivary glands in Nautilus, but intra-bulbar glands in relation with Owen's
faucial follicles have been described by Griffin (Zool. Bull. 1897, Vol. i. p. 149, fig. 4). Compare also
Joubin, L., " Reeherches sur la morphologie comparde des glandes salivaires. Glandes salivaires des C(5phalo-
podes." Arch. Zool. Kxper. (2) Tome v. Suppl. 1887, Mem. iii.
'^ There are simple valves (noted by Owen) at the bases of the branchio-cardiao trunks and a semilunar
valve at the root of the truncus arteriosus.
102—2
Fig. 8. View of proximal branches of the systemic
aorta, from above.
1. Truncus arteriosus (leaving the pericardium).
Main hepatic artery.
Gastric artery.
Kight hipatic artery.
Left hepatic artery.
Posterior proventricular artery.
Right posterior columellar artery.
Left posterior columellar artery.
Right pallio-nuchttl artery.
2.
3.
4.
5.
6.
7.
8.
9.
10. Left pallio-nuchal artery.
782
PERISTOMIAL HAEMOCOEL ; SYSTEMIC AORTA ; CEPHALIC ARTERIES.
right and left lobes of the liver, and, after a further course of about 15 mm. (in the
adult) sends ofif two important arteries, the hepatic artery on the left and the gastric
artery on the right of the main aortic trunk. It then turns definitely forwards dorsad
of the crop, giving off other branches (Text-fig. 8) until it reaches the intercoelic
diaphragm which separates the peristomial fi-om the peri-oesophageal haemocoel on the
dorsal side. Having arrived at this level, the dorsal aorta divides into the two main
Fig. 9. N. pompilhis, i . Dissection, from above, of the cephalic region to show the cephalic arteries. An
incision has been made through the nuchal membrane, the hood, and the buccal membrane. The brain-
capsule has been opened, and the median portion of the mantle behind the nuchal region has been
removed, b.c. buccal cone ; tent. int. extrabuceal tentacles (the external labial processes of Owen) ; La. labial
arteries supplying the buccal membrane and fringe ; s.m.a. superior mandibular artery ; s. r. superior retractor
muscles of the jaws (Owen); b.a. buccal artery; i.vi.a. paired inferior mandibular arteries; n.m. nuchal
membrane; a.c.a. anterior columellar artery; a.p.a. anterior proveutricular artery; /. ala infundibuli;
m.c. free mantle-edge; c.e. cut edge of mantle; p.n.a. pallio-nucbal artery; d.a. dorsal aorta; cer. brain
with cerebral arteries ; ft. cut edge of hood.
N.B. — Apart from the cerebral arteries, all the anteriorly directed branches of the innominate arteries pass
below the cerebral capsule. [From Quart. J. Micr. Sci. Vol. 39, 1896.]
cephalic arteries which I have formerly called innominate arteries. These are the feeders
of the peristomial haemocoel.
PERISTOMIAL HAEMOCOEL ; SYSTEMIC AORTA; CEPHALIC ARTERIES. 783
After giving off branches to the buccal mass and to the shell-muscle, the main
vessel on each side passes below the cerebral capsule, and follows the inner concave
border of the coronal (pedal) ganglion (PI. LXXXIII. fig. 28). Having arrived at the end
of the ganglionic swelling, at the point where the latter passes into the comparatively
slender pedal commissure, the vessel passes below or behind the ganglion, and then bends
sharply backwards and upwards, sending branches to the tentacles (PL LXXXII. fig. 5).
Although too much stress need not be laid upon the vagaries of blood-vessels, yet
I think that this recurrent course of the main tentacular arteries may be regarded as
an indication of an important transposition of parts or change of topography which has
taken place during the evolution of the nautiline plan of organisation.
At the point of flexure of the tentacular artery on each side a large branch is given
off which traverses the cartilage and is distributed to the alae infundibuli.
In the peristomial fundus there are symmetrically placed apertures leading into
various lacunae in the massive cephalopodiura ; there is one particularly large sinus
surrounding the lower portion of the spadix in the male (PL LXXXII. fig. 5). The
intrabulbar sinuses are of great complexity and I do not propose to analyse them. They
are all connected by more or less wide fontanelles with each other and with the
peristomial haemocoel.
With regard to the stomatogastric nerves I have nothing to add to Mr Graham
Kerr's account, except that the nerves are more deep-seated than his description might
lead one to expect. They are below the sheet-like protractor muscle and below all the
retractors, and hence it is hardly apposite to speak of them as Ijing "just below the skin"
(PI. LXXXIII. fig. 29).
The great peristomial haemocoel which is such a remarkable feature in the haemal
system of Nautilus communicates \vith the endochondral sinus by two channels which
lie between the dorsal protuberances (arcus cartilaginis) of the capito-pedal cartilage,
and the capsule of the pedal commissure (PL LXXXII. fig. 5).
These are the most important fontanelles in the wall of the endochondral sinus,
and I propose to name them the capito-pedal venous orifices. I have seen a delicate
membranous fold on the iimer wall of the endochondral sinus arching over the orifice
as if it were a valve to prevent the reflux of blood from the sinus to the haemocoel.
These are the capito-pedal valves, but I cannot certify as to their constant occurrence or
effective function though I deem it worth while to direct attention to them (PL LXXXIII.
fig. 14).
The blood of Nautilus when freshly dra-svn exudes as a colourless fluid which quickly
turns bluish at the edges of the fluid mass after exposure to the air, and the bluish
tinge gradually advances to the centre. When this has taken place the blood appears
of a uniform pale blue colour, which subsequently becomes darker. The blood does not
coagulate of itself if left to stand. It contains numerous amoebocytes, which I have
sometimes observed both in living and stained preparations to be united together in
small groups after the manner of a plasmodium, possibly the result of successive
nuclear divisions (PL LXXXIII. figg. 1 and 1 a). The nucleus of the amoebocytes presents
a chromatic network, while the protoplasm appears as a spongy reticulum, containing
granules which are dissolved by acetic acid.
784 REPRODUCTIVE ORGANS AND GENITAL ARTERIES.
11. Reproductive Organs and Genital Arteries.
The constitution and topography of the generative apparatus of Nautilus are well
known but I will consider them here briefly for the sake of topographical completeness,
and to point out the relations in the young, and also because I have formerly' described
the arteries which are distributed to the glands and ducts direct from the heart.
Owen (1832) stated that the ovary was supplied from the systemic aorta, while
Haller (1895) says that it receives branches from what I have called the posterior pallial
artery. It is interesting to recall these antagonistic statements concerning the vascu-
larisation of the gonad, since they were both made without apparent misgiving, and
are both incorrect. Of course the true facts appear more clearly after injection of fresh
material, and it would be quite impossible, I think, to discover the true genital arteries
by the most careful dissection of ordinary spirit-specimens.
In young individuals it is evident that the genito-intestinal ligament constitutes
the matrix of the gonad, which at its earliest appearance, in my material, is fully formed
but of small size, and lies upon and in the right face of the ligament behind the
pericardium. This region no doubt corresponds with its seat of origin in still earlier
stages, and serves to indicate the primary importance of the genito-intestinal ligament
which from an anatomical standpoint represents the true mesentery of the coelom
(PI. LXXX. fig. 20).
In later life, when the gonad has greatly increased in bulk, the anterior portion
of the genito-intestinal ligament which is inserted into the pallio-visceral ligament {i.e. the
perforated septum which separates the pericardium from the perivisceral coelom) appears
to be distinct from the rest, and has been named by Haller the genital ligament, but
it is not distinct in origin-.
A narrow offshoot from the genito-intestinal ligament is inserted into the distal wall
of the stomach and is the gastral ligament (PL LXXX. figg. 20 and 23). The proximal
end of the stomach, where the oesophagus enters it and the intestine leaves it, is closely
adherent to the wall of the peri-hepatic haemocoel (cf PI. LXXX. fig. 22).
Anatomical appearances in the young and adult are in favour of the gonad (ovary
and testis) of Nautilus having arisen as a simple pouch-like invagination of the germinal
epithelium into the substance of the primitive mesentery. There is no evidence of
secondary formation of a pouch in actual development, and I do not think that the
question of the primitive or secondary character of the gonadic pouch from a phyletic
standpoint is of great importance. What is of importance is the fact that the gonad
lies in the perivisceral coelom, and the sexual products virtually fall into the latter on
' Willey, A., "Letters from New Guinea, etc." Quart. J. Micr. Sc. Vol. 39, 1896, p. 172.
- Haller (op. cit. 1895, p. 202) says : — " Zuvorderst mochte ich mittheilen, dass ich das Genitalligament
und zum Theil auch das Genitointestinalligament fiir zwei solche Doppellamellen halte. die die
doppelte Anlage des Coloms beweisen, da sie meines Erachtens nach die raedianen Beriihrungswaude der
beiderseitigen Ciilomsacke darstellen. Hier sind somit diesbeziiglich noch sehr urspriinpliehe Verhaltnisse,
ahnlich wie bei den Placopboren, erhalten." This is an interesting conclusion, which my observations support
except that I find no necessity for qualifying or subdividing the genito-intestinal ligament. It is the one
mesentery, and in this capacity it is a morphological unit.
REPRODUCTIVE ORGANS AND GENITAL ARTERIES.
785
their way to the exterior. I say virtually, because the coelomic aperture of the gonadic
pouch becomes, in the adult, so closely approximated to the coelomic orifice of the genital
FiQ. 10. N. pompilius, s. Genital arteries from belnw. r. rectum; r.a. rectal artery. (N.B. — The rectal
arteries are very variable.) ant.p.a. anterior paltial artery; i.a. intestinal artery. (N.B. — This artery
usually passes to the right of the rectum, as shown in this figure ; but in one instance I have observed
it to pass down to the left of the rectum.) Iir.v. branchial veins; post.p.a. posterior pallial artery;
p.s.g. pear-shaped gland with its artery; a. branches of the preceding artery and of the gonaducal artery,
which supply the superjacent ligaments; gen. a. genital artery and its branches; t. testis; t.o. aperture
of testis; p.v.o. orifice of communication between the pericardial and visceral portions of body-cavity,
through which the posterior pallial artery passes ; gon.a. gonaducal artery ; v.s. vesicula semiualis.
(N.B. — This structure, the testis and pear-shaped gland are closely united to the heart by a ligament.)
N.o. Needhamian vesicle ; x. dotted line to indicate where the pallial duplicature merges into the body-
wall veutrally. [From Quart. J. Micr. Sc. Vol. 39, 1896.]
duct, often actually penetrating into the latter though separate from it, that the ova
and spermatozoa are conveyed directly from the gonad to the duct.
The main trunk of the median genital artery applies itself at first to the dorsal
side of the gonad, and extends round to the ventral side, giving off branches of greater
and of lesser importance.
In the female the ultimate ramifications of the genital artery reach the surfiicc of
the individual ova, the finer branches following but not confined to the reticulations
formed by the ridges of the follicle which project into the yolk (PL LXXXIII. fig. 17).
The arteries which traverse the surface of the ova (PI. LXXXIII. figg. 1.5 and 16), give
786
REPRODUCTIVE ORGANS AND GENITAL ARTERIES.
off minute branches, which pass deeply into the recesses of the follicular ridges and are
doubtless the means by which nutriment is conveyed to the vitellus. I noticed that
_- t^a.
gen. a.
Fig. 11. 'N. poinpilius. Fresh ovarian ovum showing the areolation produced by the folds of the egg-follicle
which project into the yolk. p. a. pellucid polar area, in the centre of which the germinal tract lies.
Fig. 12. Ovary of an injected specimen from above, o.a. neck of the gonad with orifice at its free end ;
o. ova seen through the transparent capsule of the ovary, with areolations partly drawn to show difference
in size of meshes in older (wide meshes) and younger eggs ; geii.a. dorsal branch of genital artery.
[From Quart. J. Micr. Sc. Vol. 39, 1896.]
the deep-lying branches anastomose with one another, thus forming a rete mirahile, while
the superficial branches appear not to form anastomoses.
The germinal tract appears in the centre of the clear polar area of the fresh egg
as a faint whitish spot upon the ventral aspect of the ova as they lie within the ovary
(Text-fig. 13). The older ovarian ova are rendered somewhat shapeless by mutual pressure
with a roughly oval outline, and measure 15"5 mm. in length with a maximum breadth
of 11 "5 mm. When the pressure is relieved by slitting open the ovary the ova round
.p.a.
Fio. 13. N. pompilius, ? . Reproductive organs seen from below in fresh condition, p.a. polar areas of
ovarian ova; o.a. orifice of the ovary projecting into the ostium abdominale of the oviduct; g. uterine
portion of oviduct ; v. vaginal portion of oviduct. [From Quart. J. Micr. Sc. Vol. 39, 1896.]
up, and those which are submature have an average diameter of about 10 mm. The
yolk is viscous and glutinous with a translucent brownish colour.
REPRODUCTIVE ORGANS AND GENITAL ARTERIES. 787
The follicular meshes' are close-set in young ova, becoming more open during the
later stages of maturation and finally disappearing before oviposition. The nearly ripe
ova rupture with the utmost facility, and this may account for certain appearances
which have been recorded by Keferstein (1865) and Haller (1895). Keferstein described
a large albuminiparous gland (Eiweissdriise) in connection with the ovary, while according
to Haller's interpretation {op. cit. p. 199) the ovary is gorged with free yolk in addition
to the ova. As I have described above, the vitellus receives its nutriment solely
through the vessels of the follicular membrane, and the adventitious occurrence of free
yolk can only be ascribed to the rupture of mature or submature ova and is therefore
an artefact.
In the female the short oviduct is closely adherent to the wall of the pericardium,
and its ostium abdominale lies behind and closely apposed to the pallio-\'isceral ligament.
The male organs of generation have received fresh description on the part of
Mr Graham Kerr (op. cit. 1895, p. 671), to whose work I may refer the reader, and also
to the explanation of the figures 16 — 19 on my Plate LXXX. The male duct com-
prises an external and an internal portion. The latter projects boldly into the perivisceral
coelom, where it constitutes a vestcula seminalis of special composition, consisting of two
portions separated from one another by a diaphragm perforated near its mesial border
(PI. LXXX. fig. 17). The ostium abdominale leads into the so-called accessory gland,
which contains a complex lumen in which the spermatophore is fabricated, after which it
is passed through a very small aperture into the distal thin-walled portion of the seminal
vesicle which I will call the antrum. Beyond this point the duct penetrates the integu-
ment at the level of the jjallial insertion by a narrow neck, and then dilates again to
form the so-called Xeedhamian sac which, as described by Van der Hoeven and by Kerr,
is partially subdivided into two cavities by an internal longitudinal septum. The septum
presents a free anterior border, in front of which the two halves of the cavity unite, and
over which the convoluted spermatophore rests as if upon a saddle (PI. LXXX. fig. 18).
The Needhamian sac is connected with the penis or terminal portion of the vas
deferens by a narrow neck surrounded by a sphincter. The penis itself is completely
di\'ided by an internal longitudinal septum (described and figured by Kerr) so that it
contains two lumina, of which onl}- the right is functional and lodges a spermatophore
at maturity, the left lumen ending blindly behind.
I have no information concerning the manner in which the spermatophore is con-
veyed from the penis to its position in the fossa buccalis on the dorsal side of the
buccal cone, where it is held by the two most dorsally placed tentacles of the extra-
buccal series on each side (PL LXXVII. fig. 6). The transparent membrane or cyst in
which the spermatophore is always contained when it occupies this position is probably
secreted by some portion of the spadix.
A mature male will have a spermatophore in the buccal fossa, anotlur in the
right penial lumen (which is capable of great distension), and a third in the Need-
hamian sac. I do not know how the spermatophore is conveyeil to the organ of
' The vascular folds of the inner layer of the follicle in the eggs of Sepia, noted in their superficial
aspects by Kolliker (1844), were first correctly interpreted and described in detail by Lankester, E. B.,
"Contributions to the developmental history of the MoUusca." Phil. Trans., Vol. 165, 1875, p. 39.
w. VI. 103
788 MECHANISM OF RESPIRATION, ETC.
Valenciennes in the female, and I have not been fortunate enough to find a fecundated
female among my preserved specimens'.
I have often taken males which had discharged the spermatophore, the empty and
ruptured cyst remaining behind in its usual position in the fossa buccalis. In captivity
the males sometimes become sperm-bound, that is over-mature, the spadix presenting
a congested appearance and the penial lumen and Needhamian sac turgid with sperma-
tophores, death resulting.
12. Mechanism of Respiration ; Branchiae and Osphradia ; Renal and
Pericardial Follicles.
The pallial insertion of the gills of Nautilus has been already referred to. They
are planted upon the mantle by means of a thick, fleshy peduncle which serves as
a massive buttress at the back of each gill, i.e. on the venous side of the gill
(PI. LXXXII. fig. 8). The inner surface of the gill, that is to say, the side which is
turned towards the middle line of the body, is the arterial surface, and the vessel
which lies below this surface leads directly into the corresponding hranchio-cardiac or
efferent branchial trunk. The outer surface, which is supported by the above-mentioned
buttress at the base of the gill, is the venous surface, and the vessel which lies below
the integument on this side is the continuation of the con-esponding reno-hranchial or
afferent branchial vessel which arises from the sinus venosus.
In Nautilus as in other Cephalopoda, and moreover as in the Diotocardia {Huliotis,
Pleurotomaria, etc.) and in the Acephala, the renal apparatus is intercalated into the
branchial circulation.
The structure of the gill of Nautilus strongly resembles that of the gill of the
Gastropoda, Prosobranchia, Diotocardia, Aspidobranchia. It consists of an axis or stem
bearing a double series of alternating branchial fulia or gill-plates, and only differs in
details from the gills of other Cephalopoda-. Especially does it agree with the latter,
and diverge from the prosobranchiate gill in the absence of cilia from the branchial
epithelium^ The stem of the gill of Nautilus arises by a broad but simple peduncle
from the inner surface of the mantle not far from the angle of insertion of the mantle
into the body-wall, and it undergoes no secondary concrescence either with the mantle
or with the body-wall, being free from base to apex. Each branchial lamella is at-
tached to the stem by means of a supporting membrane which presents a free border
on the dorsal (venous) side of the gill. The free borders of the basilar membranes of
' A spermatophore released from its ej-st and adhering to the organ of Valenciennes was first figured
by Mr Graham Kerr [op. cit. 189.5). Another instance has quite recently been recorded by Mr L. E. GrilBn
in a work which was received at the British Museum too late for extended and critical notice here.
Griffin, L. E., " The Anatomy of Nautilus pompilius," Mem. Ac, Washington, Vol. viii. fifth memoir, pp.
103 — 197, 11 text-figg. 17 plates, dated 1898 on cover, 1900 on title-page, received at the British (Natural
History) Museum on March '21th, 1902.
'^ Cf. Joubin, L., " Structure et developpement de la branehie de quelques Cephalopodes des cotes de
France." Arch. Zool. Exper. (2) in. 1885, pp. 75—150, PI. iv.— vi.
^ In Haliotis for example the whole surface of the branchial lamellae is ciliated (Wegmann, Arch. Zool.
Exper. (2) ii. 1884, p. 316). The gill-primordia are ciliated in Loligo according to Brooks {Anniv. Mem.
Boston Soc. Nat. Hist. 1880), but not in Sepia according to KoUiker {Entwick. d. Cephalopoden, 1844, p. 54).
MECHANISM OF RESPIRATION, ETC. 789
the branchial lamellae are strengthened by a pair of subcutaneous deeply staining
skeletal bars which appear to contain scattered cellular elements, but are not like true
cartilage, resembling rather a thickened basement-membrane. The skeletal bars may be
called the gill-bars, and" each branchial folium' contains a pair of them. The pro-xiraal
ends of the gill-bars, where they pass into the substance of the stem, are united together
in couples so as to form U-shaped prongs in such a way that the two limbs of the fork
pass into consecutive gill-plates.
Leaving out of consideration the gill-lamellae of the Acephala, a strictly comparable
skeletal apparatus has been found to exist in the prosobranchiate Gastropods and in the
dibranchiate Cephalopods.
In Haliotis, Wegmann (op. cit. 1884, p. 316) says that an attentive examination of
a branchial folium shows that its venous border is much more resistant than its arterial
border. The former is rigid and sharply defined, the latter soft and undulating, the
difference in consistency of the two borders being due to the existence, along the venous
border, of an internal skeleton consisting of a hyaline stylet which attenuates towards
the apex of the lamella.
In Sepia, Bume'' has described a series of branchial cartilages consisting of slender
rods (one to each gill-plate) standing out from the branchial gland (which occupies the
stem or septum of the gill) and stiffening the free basal edge of each supporting mem-
brane. The cartilage of which the rods are composed agrees in structure with the other
cartilages of the body, i.e. branching cells imbedded in an abundant hyaline matrix.
In Pleurotomaria each ctenidium', according to the careful description furnished by
the late Martin Woodward'', consists of a stout axial septum bearing two sets of triangular
gill-plates. The gill-plates are strengthened by a pair of supporting rods along their outer
(i.e. venous) margins. These rods resemble those which I have found in Jfautilus in being
flattened structures closely applied to the epidermis and enclosing between them portions
of the blood-space of the gill-plate. They lie along the dorso-lateral borders of the gill-
plates, and arch round from one plate to the next at the dorsal attachment of the
plates to the gill-septum.
It will be seen that Woodward's account of the gill-bars in Pleurotomaria might
be applied almost word for word to the corresponding structures in Nautilus.
The course of the blood through the gills has been described by Joubin^ who
has noted the absence of capillaries.
' An entire branchial folium consists of the branchial lamella proper and the basilar membrane by which
it is attached to the stem of the gill.
^ Bume, R. H., " On some points in the anatomy of Sepia officinalis L. II. On the presence of a series
of cartilages in the branchiae." P. Malac. Soc. London, 1899, Vol. m. pp. 53 — 56, text-figg.
" This is the term introduced by Professor Lankester for the true moUuscan gill as distinguished from
other adventitious respirator^' structures in certain Mollusca and from the gills of other animals.
* Woodward, M. F., "The anatomy of Pleurotomaria beyrichii Hilg." Quurt. J. Mirr. .S'c, Vol. -14, 1901.
^ Joubin, L., " Kecherches .sur I'appareil respiratoire des Nautiles." Rev. biol. Xord France, ii. pp. 409 — 428,
PI. VII. 1890. The branchial gland of Cephalopoda is regarded by Joubin as a blood-producing gland giving
origin to amoebocytes, therefore a kind of lymph-gland. He describes groups of cells (called in German
"Lymphheerde") in the stem and peduncle of the gill of Nautihig. I have also seen in the region of the
gills and osphradia and also in the siphuncle groups of large cells with reticular protoplasm which may be
lymph-glands.
103—2
790 MECHANISM OF RESPIRATION, ETC.
When an Octopus is observed at rest, attached by its suckers to a surface, its violent
respiratory movements are seen to depend upon the muscularity of the mantle and funnel,
the expansion of the mantle being synchronous with the contraction of the funnel, so
that while fresh sea-water is entering the mantle-cavity in order to bathe the gills, water
which has completed its work of oxygenation is being expelled through the funnel.
In the case of Nautilus the thin transparent mantle is closely applied to the inner surface
of the mouth of the shell, and takes no part in promoting the respiratory current of
water. The incurrent and excurrent streams are alike caused by the rhythmic con-
tractions of the funnel, more particularly of the alae infundihuli.
This is an interesting difference between the mechanism of respiration in Nautilus
and Dibranchs respectively, and was noted by me in 1896 {Quart. J. Micr. Sc. Vol. 39) in
correction of a statement by Moseley, who attributed breathing-movements to the mantled
The funnel of Nautilus is thus the essential organ of locomotion and the principal ac-
cessory organ of respiration, a combination of functions which should not be dissociated
from its anatomical structure and relations when its morphology is under discussion''.
It is not my intention to describe in detail the renal and pericardial follicles
which adhere respectively to the anterior and posterior walls of the afferent branchial
vessels, the former enclosed within the renal sacs and the latter projecting into the
pericardium.
The pericardial glands discharge a fluid excretory product into the pericardium,
which has been met with as a tenacious coagulum by Keferstein (1865) and Haller (1895),
but I have no record of this. The excretion of the renal glands, as is well kno%\Ti,
consists of solid concretions with concentric stratification somewhat as in starch-grains
(PL LXXVI. fig. 2). They are said to consist mainly of phosphates (Blasius, Keferstein).
When seen in bulk, filling the cavities of the renal sacs, their colour varies from white
through scarlet to deep crimson (cf. PI. LXXV. and LXXVI. fig. 1). I can suggest no
explanation of this variation in colour of the excretory products. The colour is retained
in many of my alcoholic specimens.
A dissection displaying the afferent and efferent vessels of the larger gill of the
right side is represented on PI. LXXXII. fig. 8, to the explanation of which I may refer
the reader.
Both the renal and the pericardial follicles are contractile in their entirety, the
systole of the heart synchronising with the diastole of the pericardial glands and with
the systole of the renal organs'.
With regard to the branchial sense-organs or osphradia which are highly charac-
1 Moseley, H. N., Notes by a Naturalist on H.M.S. Challenger, 2nd edit. 1892, p. 257. "On either side
the fold of mantle closing the gill cavity was to be seen rising and falling, with a regular pulsating
motion, as the animal in breathing took in the water, to be expelled by the siphon." The "fold of mantle"
is reallj' the ala infiindibuli.
- The mechanism of respiration of the Cephalopoda was dealt with in an instructive and suggestive
manner by Williams, T., " On the mechanism of aquatic respiration and on the structure of the organs of
breathing in invertebrated animals. Cephalopoda." Ann. Nat. Hist. (2) xix. 1857, pp. 193 — 201, PI. xv. He
points out the analogy between the contractile gills of Cephalopoda and the systole and diastole of the lungs
in pulmonary respiration.
^ I also observed rapid independent contractions of the renal organs.
MECHANISM OF RESPIRATION, ETC. 791
teristic of Gastropoda and Pelec}i3oda (Lamellibranchs), Nautilus affords a most interesting
link between the Cephalopoda and these two groups.
It appears that in the dibranchiate Cephalopoda there are no osphradia', and it
is probable that they have been lost in correlation with the much higher development
of the cerebro-visceral nervous system in these forms as compared with Nautilus.
There are two pairs of osphradia in Nautilus, one to each gill. The fact of the ex-
istence of branchial sense-organs in Nautilus was first made known by Lankester and
Bourne in 1883, and the jmir described by them is that which corresponds with the
lesser gill on each side. It appears as a small arcuate papilla between the bases of the
gills ; the sensory surface is below the arch (cf PL LXXXIII. figs. 6 — 8).
The second pair of osphradia appears as a quadrilobate papilla, often refeiTcd to as
the post-anal papilla, because it occurs behind the anus when the mantle is drawn back.
This compound papilla obviously consists of two bilobate papillae which have coalesced
in the middle line, though the degree of coalescence varies considerably, and occasionally
they are quite separate. In freshly captured specimens I soon became convinced of the
homodynamy^ of these papillae with the osphradial papillae of Lankester and Bourne, and
eventually recognised the fact that the deep fissure which causes the bilobate appearance
of the former corresponds with the angle made by the arch of the latter with the wall
of the mantle, and that in both cases the sensory epithelium was provided with vibratile
cilia, and was practically destitute of the gland-cells which are so abundant elsewhere
(PI. LXXXIII. figg. .3—8).
In the light of the preceding facts, more especially after the discovery of cilia on
the protected surfaces of both pairs of osphradia, I decided in my own mind that the
osphradial nature of the "post-anal papilla" was established, and I had seen traces of
small nerves branching out towards the osphradia from the main visceral nerves (cf PI.
LXXVIII. fig. 1), though I had not ascertained their mode of termination in the sensory
epithelium. Pelseneer has recently denied that the "post-anal papilla" is iimervated
from the visceral nerves. He says^: " Qu'il n'y a pas non plus de fibres nerveuses allant
de ces nerfs a la papille postanale elle-meme, et que celle-ci n'est pas un organe sensoriel,
comme le supposait Willey. L'extr^mit^ de ces nerfs visc^raux medians innerve seulement
le manteau et, chez la femelle, la glande nidamentaire."
Again on p. .58 of his work Pelseneer refers to the "Papilla postanale" in these
words: — "Ainsi qu'il a ^te dit plus haut, elle ne revolt pas de nerf important; elle ne
presente, ni sur elle ni autour d'elle, aucune region sensorielle. Son assimilation a une
' The cephalic sense-organ consisting of a " small antero-posteriorly directed ridge, which is placed below
and a little behind each eye," described by Mr W. E. Hoyle as the "osphradium" in Guimlus fabricii
{P. Zool. Soc. London, 1889, p. 125), obviously corresponds with the rhinophore of Xautilus. The osphradium
is often or usually described as an olfactory organ, but the term is limited to the branchial sense-organs, to
which it was first applied by Professor Lankester (Article " Mollusca," Encijc. ISrit.).
- This homodynamy was also indicated by Van der Hoevcn, although he was ignorant of their osphradial
nature. He says (Tr. Zool. Soc. London, Vol. iv. Part i. 1850) on p. 24 :—" Behind the auus [with reflected
mantle] there are on each side two small and depressed caruncles very similar to that mammillary eminence
or papilla we have seen at the root of the first branchia."
2 Pelseneer, P., "Kecherches morphologiques et phylog^nfitiques sur les Mollusques arohaiques." Hem. Cotir.
Ac. Belgique, t. lvii. 1899, p. 57 (Reprint).
792 MECHANISM OF RESPIRATION, ETC.
seconde paii-e d'osphradies fusionn^es [Willey] ne parait done pas soutenable." Dr Pel-
seneer makes no reference to my discovery of cilia' on the osphradia, otherwise he could
hardly have been so categorical as to the absence of a " region sensorielle."
Continued investigation of the organs by finer methods than I had previously employed
has resulted in a confirmation of my interpretation (PI. LXXXIII. figg. 3 — 8). Even
without this confirmation I should have confidently adhered to the view expressed upon
grounds of topography, homodynamy, and ciliation of protected surfaces, and I gather that
this view has found favour with Dr Ludwig Plate-.
The two paii-s of osphradia may be distinguished as the interbranchial osphradia and
the suhmedian^ osphradia respectively. The nerves which supply the former are not difficult
to find in section, and they have been figured in longitudinal section, i.e. in a section
parallel to the sagittal plane of the body, by Pelseneer (op. cit. PI. xxi. fig. 180).
Pelseneer says that the papilla is not sensory, but only the subjacent pallial epithelium,
which receives " un grand nombre de ramifications d'un filet du nerf visceral-branchial."
I am able to confirm this statement, with the reservation that I find that the sensory
epithelium is not confined to the adjacent pallial surface, but extends round the angle of
insertion of the papilla (PI. LXXXIII. fig. 7).
In the case of the submedian osphradia the entire sensory epithelium is raised above
the level of the pallial surface while maintaining its character as a sensory groove, for
protective purposes no doubt, and so it comes about that the papillae are bilobate with
the sensory surface occupying the depression between the lobes in each case. The nerves
are best seen in sections taken parallel to the long axis of the body, and in such sections,
if properly prepared, the nerves may be found to penetrate through the basement-membrane
of the epidermis, and to spread out in the substance of the latter external to the base-
ment-membrane (PL LXXXIII. figg. 3 — 8). In this way a remarkable intra-epidermal
fihi-iUar plexus is brought into existence comparable for example with the intra-epidermal
terminations of the collar nerve-roots of the Enteropneusta^ What is more to the point
however is that an exactly similar intra-epidetmal distribution of nerve-fibres in the osphradia
was described by Professor SpengeP in 1881 in the case of the primitive Lamellibranch
Area noae as well as in the Prosobranchs Haliutis and Trochus. The identity of distri-
bution of nerves in the osphradia of these Molluscs and of Naidilus may be said to be
demonstrative.
According to Professor Spengel's account, the osphradium of Area resembles that of
Haliotis and of Trochus in that from the subjacent visceral ganglion numerous nerves arise
which enter the pigmented cylinder-epithelium and distribute themselves in this "so dass
wir dasselbe eigenthtimliche Bild einen von starken Nervenfaserstrangen durchbrochenen
Ejjithelschicht vor uns haben wie dort."
' Willey, A., " The pre-ocular and post-ooular tentacles and osphradia of Nautilus." Quart. J. Micr. Sc.
Vol. 40, 1897, pp. 197—201, PI. x.
^ Plate, H. L., "Die Anatomie und Phylogenie der Chitonen." Fortsetzung Fauna Chilensis, Part C.
Zool. Jahrb., Supplement v. Bd. n. 1901, p. 561.
' The epithet "post-anal" in this connection is wrong both in substance and in fact.
•• Cf. Zool. Res., Part in. 1899, especially the case of Pttfchodera caritosa, p. 2-52.
'■ Spengel, J. W., " Die Geruchsorgane und das Nervensystem der MoUusken. Bin Beitrag zur Erkenntnis
der Einheit des MoUuskentypus." Zeitschr. wiss. Zool. Bd. 35, 1881, p. 375, Taf. sax. fig. 27.
eye; rhinophore ; otocyst. 793
In the sensory epithelium of the osphradium of Nautilus I have seen cells ■with large
vesicular nucleus and darkly staining nucleolus, bearing a great resemblance to ganglion-
cells, and as I have found them in special relation with the nerves it seems very probable
that they are in fact peripheral ganglion-cells which have retained their primitive position
in the epidermis. They are best seen in tangential sections (PI. LXXXIII. fig. 5).
13. Eve; Rhixophore; Otocy.st.
1 do not propose to dwell in detail upon these structures but to focus attention, for
the special purpose of this contribution, upon their topography. They are in fact inex-
tricably involved in the cephalopodium, and it is of the utmost importance to appreciate
their character of true cephalic sense-organs, whereas the ciliated ophthalmic tentacles
between which the eye is placed belong to the digital series.
The eye of the Nautilus is a hollow hemisphere with an opaque pigmented cornea
perforated near the centre by a small aperture which is the pupil of the eye. The size
of the pinhole aperture is apparently capable of slight accommodation to the intensity of
light as I have seen it contract from a comparatively wide orifice, 2'5 mm. in diameter,
to a mere slit with apposed margins. Passing from the aperture to the ventral border
of the eye is a supei-ficial pigmented groove somewhat resembling a choroid fissure or
raphe. Perhaps the development of the eye would throw some light upon the meaning
of this corneal raphe.
The dorsal border of the eye is convex, but the rest of the margin is produced into
a thin limbus. Sometimes the corneal raphe commences a short distance below the pupil,
and the latter then appears as an isolated subcentral orifice.
The eye faces outwards and cannot be moved in any other direction, the optic
peduncle being relatively immobile and capable only of slight protraction. The whole
aspect of the pedunculated eye somewhat resembles a sheath of a digital tentacle, and
Macdonald' actually asserted that there can be little doubt that the eye is a modified
tentacular sheath so fashioned as to become the seat of the special sense of \'ision. There
is perhaps no obvious inherent impossibility in this view, especially when regarded from
the standpoint of those who consider the digital tentacles to be special cephalic processes,
but I think it is important to dismiss the idea from the mind if it should ever present
itself
The eye of Nautilus was correctly described in respect of the pinhole aperture and
absence of dioptric apparatus by Owen (1832), and was subsequently examined in greater
detail by Hensen= (1805). Later still it was sho\\Ti by Fraisse^ to represent a primordial
type of eye which is also met with among the prosobranchiate Gastropods.
' Macdonald, J. D., " Further observations on the anatomy and physiology of Nautilus." P. R. Soc.
London, viii. 1857, pp. 380 — 382. In this connection the fact may be mentioned that the optic peduncle
does not contain an axial nerve but separate nerves arise from the ganglion and pass through the eye-stalk to
the retina.
2 Hensen, V., " Ueber das Auge einiger Cephalopoden." Zeitschr. wiss. Zool., Bd. xv. 1865. Nautilus, p. 203,
figures on Taf. xix. and xx. Later notes have been furnished by Haller, op. cit. 1895.
' Fraisse, P., "Ueber Molluskenaugen mit embrjonalem Typus. " Zeitschr, iciM. Zool. Bd. 35, pp. 461 — 477,
Taf. 25—26.
794 EYE ; RHINOPHORE ; OTOCYST.
The eye of the limpet {Patella) resembles that of Nautilus upon a very small scale
and bears a similar relation to the tentacle (allowing for changes in proportion and in
topography) to that which the eye of Nautilus does to the rhinophore. The latter
remark applies with even greater force to the much larger eye of Haliotis (cf my
PI. LXXVI. fig. 5), in which the cornea is likewise perforated, though the chamber of
the eye is filled by a vitreous body (Fraisse).
The eye of Pleurotomaria as described and figured by M. F. Woodward (op. cit.
1901) also resembles in principle the eye of Nautilus, the cornea being perforated, while
the cavity of the optic cup is only partially filled by the vitreous body.
In Nautilus the vitreous body is represented by a clear bacillary layer (Stabchen-
schicht) overlying the pigmented retinal epithelium, but it does not extend over the
inner surface of the cornea, which therefore appears as a black disc with the pupil in
the middle when seen from the inside (PL LXXXIII. fig. 25).
The extremely limited movements and accommodation of the eye of Nautilus suggest
that in spite of its size it is a photometric or photoscopic organ rather than a visual organ.
The rhinophore was discovered by Valenciennes (1841). It consists of a small papil-
liform tentacle placed below the eye, with an aperture at its base leading into a
deep pit lined by a ciliated epithelium, which is thro^vn into low folds. From the depth
of the pit and the ditficulty attending its dissection the designation olfactory labyrinth
seems to be justified (PI. LXXXI. fig. 6).
The otocyst was discovered by Macdonald (1855). It lies over but not in the
posterior cornu of the cartilage and immediately external to the coronal ganglion, so
that it has to be turned aside in order to expose the latter ft-om the outside (PL
LXXXI. fig. 6, and PL LXXXII. fig. 6). It contains in the fresh condition a milky
substance, which is so fluid that one experiences a certain difficulty in procuring some
of it for examination, so readily does it trickle away fi'om the ruptured otocyst. Under
the microscoj)e it resolves itself into a vast number of minute calcareous bodies called
otocones (PL LXXXIII. fig. 2). These were correctly described by Macdonald, but some-
times the contents of the otocyst harden in alcohol into a coagulum which has oc-
casionally given rise to the erroneous impression that the otocyst of Nautilus contains
a single otolith.
The eye and rhinophore of Nautilus apparently correspond %vith the eye and tentacle
of Haliotis or Strombus for example. Nevertheless this simple comparison has not always
been evident since Dr Pelseneer' in his work on the morphology of the arms of
Cephalopoda failed to grasp this relation, and apparently ignoring the existence of the
rhinophore^ claimed the ophthalmic tentacles of Nautilus as the representatives of the
t^-pical molluscan cephalic tentacles and stated that they are innervated by the supra-
oesophageal ganglion.
' Pelseneer, P., " Sur la valeur morphologique des bras et la composition du systeme nerveux central
des Ce'phalopodes." Arch. Biol. viii. 1888, see p. 730.
- Although he r'efers to the "nerf olfactif in a, footnote.
THE MOLLUSCAN FOOT. 795
14. The Molluscan Foot.
The foot of a Gastropod in its fullest development consists of a median sole' and
lateral appendages resembling metapleural folds.
Huxley (Phil. Trans. 1853) divided the median portion of the foot into three portions,
propodium, mesopodium, metapodium, and called the lateral portion the epipodium. He
pointed out that of all MoUusca the Heteropod genus Atlanta possesses the best de-
veloped foot-proper, and has its parts best specialised and separated, the peculiar ciliated
sucker of this pelagic animal representing the mesopodium.
Twenty years later Grenacher established a more fundamental subdivision of the
molluscan foot, retaining Huxley's epipodium, but viewing the entire median portion,
whether simple or differentiated into regions, as the protopodium", which is not to be
confused with Huxley's propodium.
Huxley's threefold subdivision of the protopodium still remains, though of subordinate
importance. An interesting example of it is afforded by the widely distributed species
Hurpa ventricosa, which I met with in the course of my travels. In this form the proto-
podium as a whole is capable of enormous extension so as to cover a relatively immense
superficial area, and the propodium is not only marked off from the rest of the foot by
a deep notch on each side, but Brock' has discovered that it possesses a special inner-
vation in the form of a very remarkable nervous reticulum. The mesopodium and meta-
podium of Harpa appear at a hasty. glance not to be differentiated from one another as
there is no operculum, but closer inspection reveals a line of division between them, and
if the hinder or caudal end of the foot of the living animal be held firmly in the hand
the metapodium is cast off and remains in the hand while the animal falls to the gi'ound.
This is an interesting example of muscular autotomy which was known and is referred to
in the Cambridge Natural History*, though the suggestion that it is effected by pressure
of the shell is erroneous (PI. LXXVI. fig. 3).
The metapodium is cut off with a clean concave anterior surface, the posterior surface
of the mesopodium after the act of autotomy being convex. There is no effusion of blood.
In a large species of Oliva which was common at Lifu, the protopodium is also
capable of great extension and will wrap itself round land-snails which are offered to
it as bait by the natives.
The sides of the protopodium may be produced into great natatory folds as in
Aplysiidae. These lateral expansions of the protopodium are called parapodia (Pelseneer)
or pteropodia (von Jhering). Whether or not these are of the nature of true epipodia
is a question which does not concern us here.
The epipodia are presented in their most typical development in certain proso-
branchiate genera, particularly in Haliotis, where they appear as a pair of deeply cleft
' The "foot-proper" of Huxley.
- Grenacher, H.. " Zur Entwickelungsgeschichte der Cephalopoden. Zugleich ein Beitrag znr Morphologic
der hiiheren MoUusken." Zeitschr. iciss. Zool., Bd. 24, 1874, see p. 465.
' Brock, J., "Zur Neurologie der Prosobranchier. " Zeitschr. whs. Zoo!., Bd. 48, 1889, pp. G7— 83, Taf. 6
and 7.
* Cooke, A. H., "Molluscs." Camhr. Nat. Hist., Vol. iii. 1895, p. 45.
w. VI. 104
796 CHANGES OF FUNCTION, ORGANS AND TOPOGRAPHY.
pleural folds bearing tentacles on their margins. Anteriorly the two borders of each
epipodium merge together, and the single tentaculiferous lobe encroaches in a very
singular manner over the dorsal side of the mouth between the latter and the cephalic
tentacles. There is also a nuchal membrane which arches over the head in front
(PI. LXXVI. figg. 4 and 5).
When the protopodium presents a Hat or plantar surface of reptation as in Haliotis
and most Gastropoda I shall speak of it as a platypodium. The platypodium is there-
fore to be defined as a plantar protojaodium.
The Cephalopoda and the prosobranchiate Gastropoda agree in so many features of
organisation that a definite relationship between them seems to be highly probable.
Perhaps the most fundamental difference lies in the absence of a plantar foot in the
Cephalopoda.
In a subsequent chapter I shall endeavour to show that the cephalopodium oi
Nautilus is of epipodial origin, while the siphonopodium represents the protopodium alone.
It is true that there is a discussion among malacologists concerning the true nature
of the epipodium, some authorities, foremost amongst whom is Dr Pelseneer, maintaining
the view of its pedal origin, whilst M. Boutan' is in favour of its pallial origin, and
calls it the " manteau inferieur." I would, however, derive the cephalopodium of Nautilus
from a primitive epipodium wholly irrespective of any morphological interpretation which
may be put upon the latter.
I may nevertheless suggest or rather repeat a suggestion which is implied in a
paragraph of Huxley's work (1853, p. 50)^ that the epipodium is a pleural fold of the
body-wall, and perhaps should be discussed independently of the foot as well as of the
mantle.
In the work of comparing Nautilus with other Mollusca, but especially with a Gas-
tropod like Haliotis, everj-thing depends upon the interpretation of pallial, pleural, and
pedal derivatives, or in other words, we have to ascertain the relations and modifications
of the typical molluscan mantle, epij)odium, and protopodium respectively.
15. CHANGE.S OF FUNCTION, ORGANS AND TOPOGRAPHY.
The doctrine of Change of Function (Functionswechsel) was, I think, first raised
to the rank of a moi-phological principle by Dr Anton Dohrn in 1875. The development
of organs by substitution or " Organwechsel " was first made the basis of a principle
by Kleinenberg in 1886. I am not aware that the phenomenon of translocation of organs
has been hitherto utilised for the purpose of establishing a principle of " Change of
Topography " which should have equal value with the other two in morphological inquiry.
The three principles may operate severally or in combination. Both change of function
' Boutan, L., "Le syst^me nerveux du Parmnphorus (Scntus) danB ses rapports avec le manteau, la coUerette
(manteau inferieur) et le pied." Rev. biol. Nord France, ii. 1890, pp. 449 — 478, PI. vni. and ix.
2 Huxley says the bilateral archetj-pe of the Mollusca corresponds with the vertebrate and articulate
archetypes. " The appendicular system of the Vertebrata and Articulata is represented by the epipodium in
the Cephalous Mollusca."
CHANGES OF FUNCTION, ORGANS AND TOPOGRAPHY. 797
and substitution of organs are often found associated \\'ith more or less radical changes
of topography.
One of the simplest examples of change of topography is to be met with in connection
with the position of the vent in Teleostean fishes, even in closely allied species. Instances
of this kind (as also in regard to the position of the pelvic fins of fishes) are exceedingly
numerous, but one of the most striking cases has been brought to my knowledge by
the courtesy of Mr G. A. Boulenger, F.R.S. In the Berycid genus Trachichthys the vent
is placed normally behind the series of abdominal scales, but in the closely allied genus
Paratrachichthys " as first observed by Giinther, the vent is far forward between the
ventral fins, in front of and not, as usual, behind the series of abdominal scales'."
I have selected this example of change of position of the vent, because the antero-
posterior axis of many animals corresponds more or less closely with what has been
called the oro-anal axis; it does so for example in the most primitive group of Mollusca,
the Amphineura (Chiton, Cliaetodernia, Xeomenia). In consequence of this circumstance
the oro-anal axis has frequently been adopted as a fixed quantity, from which all bearings
must be taken. This may be true in sjjecial cases but it is clearl}' not true for the
Cephalopoda, at least this is my opinion of the matter, and I do not regard the antero-
posterior axis as s}Tionymous with the oro-anal axis, although, as indicated above, I do
not forget that the two may coincide.
In Nautilus and the Cephalopoda generally, in consequence of the flexure of the
intestine there is no longer any question of an oro-anal axis, but there is an antero-
posterior axis which it is our business to discover.
The evidence of change of topography in Nautilus based upon anatomical conditions
is to be traced directly in the occurrence of conflicting axes, and in the existence of
recurrent nerves and arteries, and indirectly in the relations of the siphuncle. With
regard to the conflicting axes I am aware that these are visionary allies, since they
involve an imaginary interpretation of imaginary lines, but I have come to the conclusion
that what I have to saj' is not destitute of possible interest.
The capito-pedal cartilage, the skeletal basis of the body of Nautilus, possesses
a distinct longitudinal axis of its own, and this axis does not coincide with the axis
of the cephalopodium, but describes an angle with it- (PI. LXXXI. figg. 1 and 2).
It seems a possible \'iew to take that the skeletal axis may represent the primitive
longitudinal axis of the uncoiled ancestor of Nautilus, that the tilting of the cephalopodial
axis is evidence of the concrescence and translocation of the primitive epipodium (the
tentacles of Nautilus being here regarded as epipodial tentacles), and finally that the
ventral pallial flexure of Nautilus has resulted in the formation of a visceral sac or
abdomen, which lies chiefly below the level of the primitive antero-posterior axis
(PI. LXXXI. fig. 2). The lateral portions or comua of the cartilage which determine
the direction of its main axis, occur along the angle of junction of the epipodium
(cephalopodium) and protopodium (funnel).
The recurrent nerves of the posterior ophthalmic tentacles obviously indicate change
1 Boulenger, G. A., " Notes on the classification of Teleostean Fishes. 11. On the Berycidae." Ann. Nat.
Hut. (7) IX. 1902, p. 203.
' These relations are best seen in dissections of relaxed specimens.
104—2
798 . FLEXURE AND ORIENTATION.
of primitive topographical relations of parts, and perhaps a similar interpretation, although
not so certain, may be put upon the recurrent tentacular arteries (PI. LXXXII. fig. 5,
and PI. LXXXIII.''fig. 28).
With regard to the siphuncle, while it is certain that it has a physiological relation
to the air-chambers of the shell, its morphological nature is not so clear, but I think
it is a vestige, a venniform appendix of A^'autilus, the remnant of the primitive elongate
body which we must suppose the remote ancestor of Nautilus possessed. The gut and
the coelom have vanished from the siphuncle, and it is now reduced to the condition
of a vascular appendage, of vital importance to JS'autilus in a physiological sense, but
a mere vestige morphologically.
16. Flexure and Orientatiox.
When squids are watched darting through the water with their pointed hinder
extremities turned in the direction of locomotion, the possibility of there being any
question concerning the orientation of the animal, i.e. concerning the true bearings of
the antero-posterior axis, never occurs to the observer. It is left to the refinements
of morphology to raise such a question, the discussion of which is only too apt to be
profitless and uninteresting. It is however impossible to neglect it in a work like the
present.
Considered as Mollusca, the problem of the orientation of the body of Cephalopoda
depends upon and is created by two main features of organisation: — (1) the absence
of a plantar foot or platj^odium ; (2) the ventral position of the forwardly directed anal
aperture, which is associated \\*ith what Huxley (1853) called a neural flexure of the
intestine.
The current notions regarding the orientation of a Cephalopod follow the lines laid
do^vn b}- Huxley, according to whom the visceral sac represents a dorsal hump or
abdomen while the arms represent a ventral foot, so that the morphological position of
the animal is taken to be that which is sometimes assumed for example by Octopus
(cf also my PL LXXVII. fig. 3) in which the arms are directed downwards, i.e. towards
the substratum and the visceral sac upwards. This is also the position which is usually
postulated for Nautilus on the tacit assumption that Nautilus in spite of its possession
of many primitive anatomical characters, affords no fresh clue in this regard.
Another view with which I find mj-self, under certain reservations, in substantial
agreement, has been started in recent years by Dr B. Haller. I may quote the following
paragi'aph fi-om this author's work on Nautilus (op. cit. 1895, p. 191): " Bezliglich der
Orientirung mochte ich vorausschicken, dass ich den Nautilus, vde uberhaupt die Cephalo-
poden (wie ich dies ausflihrlicher schon mitgetheilt habe' ) aus einem chitonartigen
Mollusken so hervorgegangen mir vorstelle, dass letzterer mit seinem hinteren Korper-
abschnitt sich von hinten nach vom und somit nach ventral warts bog, wodurch
der After nach vom zu gelangte Vom ist der Kopf, hinten der Sipho."
' Haller, B., " Studien iiber docoglosse und rhipidoglosse Prosobranchier, etc." Leipzig, 189-t, p. 149. The
"Sipho" means of course the siphuucle.
FLEXURE AND ORIENTATION, 799
In the course of the following pages I shall give my ovm reasons for regarding
the cephalopodium of Nautilus as being anterior in position and (ftir from being ventral)
having a marked dorsal inclination by which, as mentioned above, its main axis as
determined by the direction of the sheaths of the digital tentacles has become tilted
at an angle with the skeletal axis (PI. LXXXI. figg. 1 and 2)'.
With regard to the flexure or coiling of the external, multilocular, involuted shell
of Nautilus, it was noted by Huxley (1853, p. 49) that the direction in which it is
wound is the same as that in which the intestine is bent, so that if the coils of the
shell be imagined free as they are in Spirtda, the convexity is dorsal and the concavity
ventral. This method of coiling has been called exogastric by Owen-, while the shell
of Spirula which is coiled in the reverse direction is endogastric. Spirula is a coiled
Belemnite (Valenciennes), Nautilus a coiled Orthoceras (Owen), and Ammonites a coiled
Baculites (Owen).
The shell of Nautilus is external and protective, it is an involute cone of great
length, and its length incxeases pari passu with the growth of the animal until the
latter reaches its specific cubic capacity or limit of growth. The deposition of septa
and secretion of gas convert the shell into a hydrostatic apparatus.
Owen regarded the straight chambered shells of fossil Cephalopoda as having been
produced by the progressive unwinding of a coiled shell and the evidence seems to point
in this direction.
If this be so, then the origin of the shell is lost in the remotest antiquity, and
it is useless to discuss the question whether the arthrocochlid (von Jhering) shell-system
of the Polyplacophora (Chiton) has any possible relation to the septate shell of the
primitive Cephalopoda.
The actual fiicies of Nautilus is determined by its visceral flexure, and by its
pedal and epipodial tojDography, and this is therefore the place to refer to a question
which has been raised by Dr Plate ^ namely, as to whether the siphonopodium (Schwimm-
fuss) of Cephalopoda is derived from a platj-podium or vice versa. It seems to me that
the direct answer to this question depends entirely upon one's individual proclivities.
The cleft funnel of Nautilus conveys the impression of a platypodium, with the margins
folded over each other in a manner recalling that in which the large extensile f(3ot
of certain Gastropods (Harpa, Oliva, etc.) is capable of wrapping itself round foreign
objects.
If we imagine the body of Nautilus to be straightened out and the anus carried
back to its primitive terminal position, the dimensions of the primitive platypodium,
extending from end to end of the body, may appear apjJalling. It is however quite
conceivable that the enormous bulk of the Cephalopods, as a whole, was correlated with
the achievement of the peculiar relations of the animals to their shells, and therefore
came into being subsequently to the evolution of the plan of composition of the
Cephalopod organisation.
■ I may also refer the reader to the two preceding chapters.
- Owen, E., " On the relative positions to their constructors of the chambered shells of Cephalopods."
P. Zonl. Soc. London, 1878, pp. 955—975, PI. 40.
s Plate, H. L., op. cit. 1901, p. 559.
800 MORPHOLOGY OF THE TENTACLES OF NAUTILUS.
The siphonopodium seems to be a more highly specialised organ than the platy-
podium, and for this direct reason in combination with collateral considerations, it is
probably more useful at present to accept the dogma that the foot of the molluscan
archetype was a platypodium.
17. Morphology of the Tentacles of Nautilus.
The two theories of the special nature of the tentacles of Nautilus, considered
merely from a cephalopodan standpoint, have been referred to in a former chapter, and
may be tabulated in chronological order as follows: —
1. Acetabular Theory of Valencieimes (1841), according to which a tentacle of Nau-
tilus, consisting of the sheath and the cirrus, is homologous with a sucker of a Dibranch
consisting of cupule and caruncle.
2. Branchial Theory of Owen (1843), according to which a tentacle of Nautilus is
homologous with an arm of a Dibranch.
Both of these theories have received authoritative support, and neither of them is to
be lightly rejected, since neither is capable of such final jjroof as would satisfy the higher
criticism.
The independent innervation of the individual tentacles of Nautilus, the demonstration
of their definite suctorial ridges by which they are enabled to adhere to surfaces with great
tenacity, and the virtual identity of their internal structure with that of a true arm,
weigh so heavily, to my mind, on the side of the Branchial Theory, that I must follow
Owen in admitting their equivalence with the arms of the Dibranchs just as the multi-
tudinous legs of a Myriapod are individually equivalent to the legs of a Hexapod insect.
It was further suggested by Owen that the inner whorl of the cephalopodium of Nautilus
may be represented in the decapod Calamaries and Cuttle-fishes by the two long re-
tractile prehensile arms or anchors, which are provided at the base with a kind of sheath.
It is worth repeating that the suctorial ridges of Nautilus actually tend to become
semilunar or half-cujJ-shaped towards the apices of the outermost tentacles, a feature
which is especially noticeable in the ophthalmic tentacles. Nevertheless it is quite possible
that a single ridge does not corresjDond with a single sucker, but rather with a trans-
verse row of suckers such as occurs in Gonatus, where the arms carry four series of
suckers'. A curious analogy is afforded by the second and third tentacles of the spadix
in Nautilus, the transverse costae of the former being represented by the transverse rows
of alveoli in the latter (cf PI. LXXIX. fig. 9).
In support of the Acetabular Theory of Valenciennes it is of great historical interest
to know that the case of Girroteuthis was quoted by Johannes Miiller at a meeting of
the Berlin Academy on the occasion of the communication by Humboldt of an abstract
of the results achieved by Valenciennes in his work upon the anatomy of Nautilus^.
Johannes Miiller is reported to have said that the Girroteuthis of Eschricht has arms which
1 Cf. Hoyle, W. E., op. cit. P. Zool. Soc. London, 1889.
' Bericht Akad. Berlin, 1841. Bemerkungen des Hrn. Prof. J. Miiller, p. 58. Translated in Ann. Nat.
Hist. Vol. VII. 1841, pp. 241—245.
MORPHOLOGY OF THE TENTACLES OF NAUTILUS. 801
are not furnished with suckers but with delicate filiform tentacula, but the fact is that
there is a median row of suckers as in Eledone, though the suckers are reduced, and
alternating with them are 26 pairs of cirri on each arm'.
When we come to consider the broader interpretation of the arms of Cephalopoda
from a pan-molluscan standpoint we are again confronted with two rival theories which
may be defined as follows :^
1. Pedal Theory of Huxley' (1853), according to which the arms of Cephalopoda are
derivatives of the primitive moUuscan foot.
2. Cephalic Theory of Leuckart (1848) and von Jhering (1877), according to which
they are special appendages and derivatives of the primitive head.
The most complete exposition of the Pedal Theory is contained in the memoir by
Pelseneer^, while the Cephalic Theory has been strongly advocated by Grobben*, and more
recently by Kerr'.
Huxley, as is well known, regarded the arms of Cephalopoda as representing the
protopodium and the funnel the epipodium ^ " The formation of an abdomen," he said,
" with a peculiar development of the margins of the foot into elongated processes, and
with cohesion of the posterior epipodial lobes, gives us the Cephalopodan subtj-pe."
From this quotation it is clear that Huxley considered the arms as marginal appendages,
which is in itself an important conclusion. His reason for identifying them with the
protopodium may possibly be looked for in the bionomical fact that Octopus, for example,
actually does crawl about fi-om place to place by means of its arms, although it can
s\vim vigorously on occasion. The identification of the funnel with the epipodium may
be due to the fact of the cleft funnel of Nautilus, the flaps of which may be likened
to pteropodial lobes.
Grenacher {Zeitschr. wiss. Zool., Bd. 24, 1874) homologised the aims with the velum
of Gastropod embryos, but adopted Huxley's interpretation of the funnel on account of
the development of the latter from paired primordia.
The special reasons for the view that the funnel is an epipodial derivative are there-
fore based upon facts of bionomics, comparative anatomy, and embryolog)', but I believe
that these are facts which mask the truth in this particular case. The epipodium as
the name implies lies dorsad of the protopodium, while the siphonopodium of Cephalopods
is ventral in position, and although great allowances may be made for changes of topo-
1 Eschricht, Cirroteuthis mUlleri; Nona Acta Ac. German, t. 18, 1836, p. 627, tabb. 46 — 48. "Octopus
snctoriis minimis unam seriem in quovis brachio formantibus ; braohiis cirratis et cum merabrana natatoria
usque ad apicem fere connatis."
^ Huxley, T. H., " On the morphology of the cephalous Mollusca as illustrated by the anatomy of certain
Heteropoda and Pteropoda collected duriug the voyage of H.M.S. Rattlesnake in 1846 — 50." Phil. Trans.
Vol. 143, 1853, pp. 29—65.
* Pelseneer, P., " Sur la valeur morphologique des bras des C^phalopodes." Arch. Biol., viii. p. 723,
18H8; see also Dr Pelseueer's "Report on the Pteropoda." Chall. Rep., Part 60, Vol. xix. 1888. Anatomy
(Part in. of Report) Chapter ii. "Are the Pteropoda Cephalopoda?" p. 60.
■• Grobben, C, " Morphologische Studien iiber Cephalopoden." Arb. Inst. U'ien, v. 1884, see p. 222
(p. 44 of Separat. Abd.). Also Grobben, C, " Zur Kenntniss der Morphologie und der Verwaudtschaftsverhiiltnisse
der Cephalopoden." Arb. Inst. Wien, vii. 1886, pp. 60—82.
° Kerr, J. G., op. cit. P. Zool. Sac. London, 1895, see p. 678.
6 Huxley, T. H., op. cit. 1853, PI. v. fig. 5, p. 51.
802 MORPHOLOGY OF THE TENTACLES OF NAUTILUS.
graphy it is difficult to admit the probability of such a complete situs inversus, nor is it
necessary in order to co-ordinate the facts.
The only author who has referred to the possibility of the epipodial origin of the
cephalopodium, so far as I know, is Professor Brooks', and he adopts a somewhat pessi-
mistic attitude in the matter. Professor Brooks commences his remarks by referring to
what may be defined as the Velar Theory of Loven (1848) and Grenacher (1874), by
whom the arms of Cephalopoda were compared with the velum of other moUuscan em-
bryos, but it cannot be said that this theory has enjoyed a great measure of success, and
Brooks has found that in Loligo, at the stage at which the ingrowth of the stomodoeum
takes place, the rudiment of a true velum appears on each side of the mouth in the
form of a faintly marked, undulating, ciliated band arising near the comers of the
mouth, and extending laterally to the eye-stalk, crossing the outer ventral edge of the
latter. The position of this line, its relation to the mouth and the eye-stalk, and the
presence of cilia upon it, are all indications that it represents a true velum.
Professor Brooks adds that " the siphon originates as two pairs of folds - . . . . and
if we regard these four folds as homologous with the epipodial folds of a Gastropod,
the arms must be regarded as independently acquired structures. If we regard the
arms as modifications of the epipodial folds we must consider the four siphon folds
as independently acquired structures^ and as we have nothing whatever to furnish us
with a test, nothing seems to be gained by the uncertain homologj' of either the
arms or the siphon, with any part of the body of a t}'pical Gastropod."
On the other hand I think it will be no mean gain for morphology if we can
secure the recognition of the epipodial nature of the arms and the protopodial nature
of the funnel of Cephalopoda, and the longer I meditate upon the subject the safer do
these homologies appear to my mind. It is, as everyone knows, a difficult matter to
adduce a clinching argument in favour of a theoretical conclusion, and if the cumu-
lative weight of what has been brought forward in the preceding pages is not suffi-
cient to turn the scale, I fear it will be impossible to say more with advantage.
The following is a summary of the special grounds upon which I base my Epi-
podial Theory : —
1. Tentacles of Nautilus regarded as marginal appendages.
2. Sequence of the tentacles of Nautilus and of their nerves.
3. Topography of cephalopodium and siphonopodium, the dorsal position of the
former, the ventral position of the latter, and the deep longitudinal pleural raphe which
separates them.
4. Comparison with Haliotis where we find in schematic disposition, mantle, epi-
' Brooks, W. K., "The development of the squid, LoUao pealii (Lesueur)." Anniv. Mem. Boston Soc. 1880,
22 pp., 3 Plates.
- These folds clearly correspond with the anterior crura infundibuU and the posterior alae infundibuli of
the funnel of Nautilus.
' This is equivalent to saying, what I believe to be true, namely, that the method of development of
the funnel from paired primordia, is a cenogenetic feature in the embryos of Cephalopoda which may be
correlated with the special adaptation of the protopodium which has resulted in the formation of the funnel
and with the abundance of yolk in the eggs of Cephalopoda, in which respect Xautilui surpasses all other
known genera.
MORPHOLOGY OF THE TENTACLES OF NAUTILUS.
803
podium and plat3rpodium, these structures being represented in Nautilus by the mantle,
cephalopodium and siphonopodium respectively.
5. The recognition of a dorsal and a ventral symphysis or lines of concrescence
in the cephalopodium of Nautilus.
6. The relation of the cephalic sense-organs to the cephalopodium of Nautilus and
their homology with the corresponding organs in Gastropoda.
7. The presence of an outer and an inner whorl of tentacles in the cephalopodium
of Nautilus in apparent correspondence with the double nature of the tentaculiferous
epipodium of Haliotis.
8. The presence of the funnel-organ which according to Jatta represents a pedal gland.
9. The position, form and relations of the capito-pedal cartilage of Nautilus.
A word may be added by way of justification for the attempt to construct the above
theory in the absence of embryological evidence. The enormous size of the vitellus of
the egg of Nautilus renders it highly probable that the development of the tentacles and
funnel would be direct, it would take place in situ, and the topography of the adult would
be maintained in the embryo, all special embryonic events, although of the greatest
possible interest in themselves, being of a cenogenetic nature'. I have actually observed
that the inner whorl of the cephalopodium arises in a perfectly direct manner (PI. LXXX.
figg. 1 and 2).
The following tabular view of the principal theories relating to the arms of Cephalo-
poda will at once illustrate the complexity, interest, and importance of the subject.
Author
Cephalopodium
Siphonopodium
Valve
Funuel-organ
Protopodium
Huxley
Protopodium
Velum
Epipodium
Epipodium
Arms
Grenacher
Wanting
von Jhering...
Cephalic tentacles
Pteropodium
Protopodium
Valve
Lankester
Propodium
Mesopodium
Metapodium
Arms + Funnel + Valve
Neomorph
Cephalic captaoula
(as in Dentalitim)
Protopodium
Neomorph
EpijKjdium
Epipodium
Neomorph 2
Protopodium
Yolk-sac
Grobben
Pelseneer
Valve
Arms
Jatta
Cephalic processes
Epipodium
Protopodium
Protopodium
Pedal gland
Arms + Funnel
Funnel
Kerr
Willey
Neomorjih^
Pedal gland
Funnel
' This does not refer to the development of the shell and siphuncle, which is likely to afford many
instructive data. Even with regard to the cephalopodium important facts might be ascertained regarding the
concrescence of the pleural folds.
^ In Loligo the valve develops at a late stage as an outgrowth from the inner wall of the funnel (Brooks).
When seen from the inside of the funnel with the flaps of the latter turned aside (cf. my PI. LXXXIII.
fig. 27) the free border of the valve is seen to be concentric with the anterior border of the funnel. Between
w. VI. 105
804 DIPLOMERISM OF NAUTILUS.
Should the foregoing inferences appeal to malacologists in general, an important de-
duction relating to the homologies of the nervous system remains for consideration. It
would follow that the coronal' ganglion and commissure of Nautilus and their outgoing
nerves are equivalent to the pleuro-pedal system of prosobranchiate Gastropods, and that
here, as in Nautilus, the central and peripheral nervous apparatus is composed of two
principal systems in close union with one another by means of connectives, but otherwise
distinct, namely, the cerebro-visceral system and the pleuro-pedal system.
18. DiPLOMERISM OF NaUTILUS.
There are indications of two distinct metameres in the trunk of Nautilus, and the
demonstration of the osphradial nature of the " post-anal papillae " adds another link to
the chain.
0\ving to the visceral flexure of the mantle which involves renal organs, gills, rectum,
etc., in other words, owing to topographical readjustments occasioned by life within a
shelP, the segments do not occur entirely in metameric succession, but the morphologically
anterior segment lies ectad and dorsad of the morphologically posterior segment, and even
behind the latter. A representation of the two segments is contained in the figure on
PI. LXXV., which shows the pallio-visceral region of Nautilus seen from below through
the transparent mantle during life, with natural colours.
In order to avoid repetition of the terms " anterior " and " posterior " now in a mor-
phological, now in a topographical sense, I will call the morphologically anterior segment
the outer or microbranchiate segment, since it comprises the lesser gill ; the morphologi-
cally posterior segment is the macrohranchiate or submedian segment.
I. Microbranchiate segment contains on each side of the body: —
1. The smaller branchia.
2. A reno-branchial vessel, to which are appended
3. The outer pericardial gland, and
4. The outer renal organ which lies in
0. The outer renal chamber which opens to the exterior by
6. The outer renal orifice.
7. The posterior branchio-cardiac vessel.
8. Outer pericardial ligament (PI. LXXXII. figg. 7 and 8).
9. Interbranchial (outer) osphradium.
10. Outer \'iscero-branchial nerve.
11. Outer viscero-osphradial nerve.
12. Generative orifice on right side and orifice of pyriform body on the left.
the valve and the dorsal wall of the funnel there is a deep angle or groove and it seems possible to compare
this with the anterior sulcm which characterises the border of the propodium in Harpa (PI. LXXVI. fig. 3),
PleuTotomana (Woodward) and other prosobranchiate Gastropods. This sulcus is the "glande pedieuse
ant^rieure" or " sillon du bord anterieur du pied" (Pelseneer, Introd. a I'etwie des moUusques, 1894, p. 52).
' Generally called " pedal."
- Or accompanying it.
AFFINITIES. 805
II. Macrobranchiate segment contains on each side : —
1. The greater branchia.
2. A reno-branchial vessel, to which are appended
3. The inner pericardial gland, and
4. The inner renal organ which lies in
5. The inner or submedian renal chamber which opens to the exterior by
6. Its owTi renal orifice.
7. The anterior branchio-cardiac vessel.
8. The inner (anterior) pericardial ligament.
9. The inner or submedian osphradium.
10. Inner viscero-branchial nerve.
11. Inner viscero-osphradial nerve.
12. The viscero-pericardial orifice.
The outer visceral nerve supplies both of the gills and the interbranchial osphradium.
The inner visceral nerve supplies the mantle (nidamental gland in female) and the sub-
median osphradium.
The heart of Nautilus has undergone a semi-rotation by which the origin of the
systemic aorta is can-ied backwards and the positions of the branchio-cardiac vessels perma-
nently inverted, the anterior pair draining the greater (morphologically posterior) branchia,
while the posterior pair is associated with the lesser (morphologically anterior) branchia.
The diplomerism breaks down in a somewhat puzzling fashion when we come to con-
sider the coelom, since the pericardium and the perivisceral coelom can hardly be regarded
as homodynamous subdivisions of the secondary body-cavity. There is therefore no true
coelomic metamerism in Nautilus in spite of the existence of metameric abdominal pores,
namely, the generative and viscero-pericardial orifices, the latter at any rate taking the
place of the nephrostomes of other Cephalopoda'.
An interesting, analogous, physiological interrelation between nephrostomes and ab-
dominal pores has been established by Bles'' in fishes.
19. Affinities.
I do not propose to undertake an exhaustive discussion of the affinities of Nautilus
beyond what has been already said. With regard to the relationship of Nautilus and
Cephalopoda in general to the Amphineura, the most primitive of existing MoUusca,
in which the antero-posterior axis coincides with the oro-anal axis, I will refer the reader
to the works of Kerr, Haller, and Plate to which I have already referred'.
> Cf. Jhering, H. v., Zeitschr. wiss. Zool., Bd. 35, 1881. Grobben, C, Arh. Imt. U'icn, v. 1884 and
VII. 1886. Kerr, J. G., P. Zool. Soc. London, 1895. Ziegler, H. E., " tjber den derzeitigen Stand der Ciilomfrage."
Verh. Deutsch. Zool. Ges. 1898, p. 14.
' Bles, E. J., "Abdominal pores and nephrostomes in fishes." J. Ariat. I'liysiol., Vol. xxxii. p. 484; also
P. R. Soc. London, Vol. 62, 1898, p. 232.
' See further Kerr, J. G., " Phylogeuetic relationship between Amphineura and Cephalopoda." Zool. Anz.,
XXIV. 1901, p. 437.
105—2
806 AFFINITIES.
It is difficult to know how to regard the Aplacophora {Neomenia, Chaetoderma), but
as for Chiton itself, the constant octomerism of the shells seems to indicate that it is
a highly finished type which has retained primitive relations of main axis and bilateral
symmetry.
Kerr holds that the Amphineura are the nearest living allies of the Cephalopoda,
and he gives instructive diagrams of their coelomic properties. In respect of the mutual
relations between coelom and haemocoel. Nautilus is particularly interesting, and in respect
of its possession of a spacious perivisceral coelom incompletely divided into two halves
by a median mesentery, the genito-intestinal ligament, Nautilus seems to present a more
primitive organisation than any other existing mollusc.
Grobben (1884-1886) was of opinion that the Scaphopoda {Dentalium) were the
nearest living allies of Cephalopoda, but this view has not been sustained by Simroth
in his most recent treatise on the former group in Bronn's Thierreick.
For my part I have found, during a somewhat limited experience of malacological
matters, that a comparison of Nautilus with the prosobranchiate Gastropods, more especially
with Haliotis, seems to throw considerable light upon the organisation of the former genus
and indirectly of the dibranchiate Cephalopods.
The torsion of the palho-visceral region of Haliotis with the correlated streptonexirism
of the visceral commissure, in spite of the profound effect it has had upon the facies of the
animal, may be regarded from a morphological standpoint as an incidental phenomenon
equally with the pallio-visceral flexure of Nautilus.
If abstraction be made of the torsion of Haliotis and the flexure of Nautilus the
general resemblance between the two genera can be more readily conceived, and the
relation of the Cephalopod to the Gastropod would appear in the same light as that
which a Hexapod insect bears to a MjTiapod, or a Decapod Crustacean to a Phyllopod.
If we take into consideration the analogous phenomena of cephalisation in different
groups of coelomate animals, it becomes evident that the general tendency is to evolve a
composite head and a concentrated central nervous system. In this sense the cephalopodium
of the most highly organised group of MoUusca is analogous with the cephalothorax of
a crab or a lobster. There is thus undoubtedly a prima facie case for the composite
nature of the cephalopodium, tending to exclude the idea that this structure can be
merely a derivative of the head without the co-operation of any part of the trunk.
I have endeavoured to show that Nautilus presents certain features of organisation
which have become obliterated in the Dibranchs, but which are of crucial significance
in regard to the interpretation of the organisation of Cephalopoda, and this without
regard to the diplomerism and perivisceral coelom of Nautilus. Chief among these
nautiline characteristics, I reckon the remarkable tilting of the cephalopodial axis at an
angle to the skeletal axis (PI. LXXXI. fig. 1). If this is a sound description of facts
it must be of importance in itself, and especially because no such conflicting axes can be
observed in the Dibranchs. I interpret it as meaning that a tentaculiferous epipodium
has coalesced with the head and grown round the buccal cone both above and below,
while the protopodium has suffered a greatly inferior modification of structure, simply
having its margins folded over to form a fuimel.
In the commonly accepted orientation of Cephalopoda, no difference is recog;nised
AFFINITIES.
807
between Nautilus and Dibranchs, and Nautilus is not specially consulted in the matter,
and yet I think it is alone a competent witness.
The similarities between Nautilus and a diotocard Prosobranch {Haliotis, Fissurella,
Pleurotomaria), whether of the nature of affinity or of convergence, include the following
points.
System. Tetrabranchiata.
lAoiia inferior s. pallialis.
\ Aorta superior s. systemica.
Circulatory. -.Intestinal branch of pallial artery.
Reno-branchial system.
Nervous.
Sensory.
Motor.
Branchial.
Pallial.
Reproductive.
Digestive.
Peristomial haemocoel.
rPleuro-pedal collar.
J. Intra-epidermal fibrillar
[ plexus of osphradium.
Eye with perforated cornea.
[ Cephalopodium.
( Siphonopodium.
(Branchial folia biserial.
(Branchial skeleton.
Nidamental gland.
Dioecism.
(Pyloric coecum.
Ciliated intestinal epithelium.
Absence of extrabulbar salivary glands.
Prosobranchiata.
Arteria pallialis.
Truncus arteriosus communis.
Rectal branch of pallial artery.
Similar.
Cephalic sinus.
Pleuro-pedal cords.
Same.
Same.
Epipodium.
Platypodium.
Same.
Same.
Hypobranchial gland (Haller, 1894).
Same.
Similar.
Same.
Same.
With regard to the relationship of Nautilus to the other Cephalopoda opinions will
always differ as to details, but the main issues are clearly defined. In respect that it
is a Cephalopod, the external shell of Nautilus is probably primitive as compared with
the internal shell of Spirula and the other Dibranchs'.
Upon the primitive characters of the digestive system, the paired liver, structure
of pyloric coecum, etc., Haller (1895) may be consulted. Brock tabulated upwards of
a dozen different combinations and correlations of oviducts and nidamental glands, and
pointed out that the slit-like orifices of the renal sacs of Nautilus are of primitive
significance, inasmuch as this kind of renal orifice is again met with in all Oigopsid
cuttle-fishes, and in the more primitive forms of the other subdivisions, whereas in the
higher Myopsidae and Octopoda a fleshy renal papilla has developed^. He thinks that
the Octopoda are the most highly specialised Cephalopoda (as illustrated in the structure
of the central nervous system, the eye which can be closed by eyelids, the atrophy of
' On vestigial shells of Cephalopoda see Appellof, A., " Uber das Vorkommen inuerer Schalen bei den
achtarmigen Cephalopoden (Octopoda)." Bergens Mus. Aarhog. 1898, No. 12.
^ Brock, J., " Studien iiber die Verwandtscliaftsverhaltnisse der dibranchiateu Cephalopoden." S. B. Soc.
Erlangen, 1879, Heft 11, pp. 114—141. Also Morph. Jahrb., vi. 1880, p. 185.
808 FOOD ; MIGRATION ; PROPAGATION.
the shell"), but that they retain many primitive features such as the crop, the septum
between the renal sacs, the paired oviducts.
20. Food; Migration; Propagation.
The specimen of Nautilus which wiis dissected by Owen contained fragments of
triturated Crustacea in its stomach and crop, and this would seem to constitute its staple
food-supply since prawns and crabs abound on the nautiline terrains.
As mentioned in a previous chapter any kind of animal bait will tempt Nautilus,
and after a full meal the crop is found to be gorged to repletion.
In order to be in a position to form a just estimate of the value of the various
accounts and legends concerning the habits of Nautilus several facts require special
consideration. In the firet place Owen drew attention to the circumstance that whereas
two kinds of Nautili are referred to by Aristotle, only one kind is mentioned by Pliny.
In his Thesaurus Conchyliorum, Sowerby (1855) took care to note particularly that
the Pearly Nautilus is not the Nautilus of Pliny, which is Argonauta or the Paper
Nautilus, so-called on account of the thin, white, delicate structure of its peculiar shell.
No doubt many of the fantastic ideas concerning our Nautilus, more especially such
as possess the popular mind, owe their origin to a simple confusion between the Pearly
or, as we may picturesquely term it, Aristotle's Nautilus and the Paper or Pliny's Nautilus.
That this is the case is indicated in a convincing manner by the identity of some of
the myths which have been related in connection with both species'.
It is also desirable to remember that Nautilus obviously draws its supplies of food
from the bottom of the sea, it is a ground-feeder, while Argonauta, according to available
evidence, is a pelagic feeder.
When Nautilus has been taken, as a gi-eat rarity, at the surface of the sea, it has
generally, if not always, been found that the specimen was in a more or less moribund
condition. At the same time, with its known faculty for swimming and migration in
some places into quite shallow water a few fathoms only in depth, it is quite conceivable
that an individual specimen might occasionally wander away from its home and arrive
at the surface, but there is no evidence that this is a regular practice.
In his " Notes by a Naturalist on H.M.S. Challenger," Moseley also gives it as his
opinion that " it is probably a mistake to suppose that it ever comes to the surface
voluntai-ily to s\vim about."
Moreover only single individuals have ever been seen at the surface and then in the
daytime'-, but the evidence of the traps goes to show that Nautilus is a gregarious
animal and nocturnal in its habits. It repairs in shoals at night to its shrimping
grounds, but I suspect that it breeds in deep water or in inaccessible submarine gullies.
1 On the subject of the legendary accounts of the habits of Argonauta, the following interesting paper
may be consulted ; — " Observation d'un Argonaute de la Mediterranee," by H. de Lacaze-Duthiers, Arch. Zoo!.
Exper. (2) x. 1892, pp. 37 — 56, with figure of the animal in swimming attitude on p. 41.
^ E.g. Saville-Kent, W., "Preliminary observations in connection with the surveying cruise of
H.M.S. Myrmidon at Port Darwin and Cambridge Gulf." P. It. Soc. Queensland, vi. p. 229, 1888.
FOOD ; MIGRATION ; PROPAGATION. 809
In his account of the specimen of Spirula obtained by Sir Edward Belcher during
the voyage of H.M.S. Samarang 1843 — 1846, Owen (Zoology of the " Saniarang," London,
1850, Mollusca, p. 6) called attention to the fact that the two known genera of ex-
isting polythalaraous Cephalopods were noted for the extreme rarity of the entii-e animal
as compared with the frequency of the shells in collections of Natural History. In the
remaining Report on the Mollusca of the " Samarang," Messrs Arthur Adams and Lovell
Reeve recorded the fact that no living Nautilus was seen during the whole course of the
expedition, although it was assiduously sought for.
There is not much to be added here concerning the propagation of Nautilus. Some
animals breed freely in confinement, others become egg-bound, and others again lay sterile
eggs, as Nautilus did in my cages. I obtained some evidence that the captivity had the
effect of interfering with the normal intercourse of the sexes. On one occasion in parti-
cular I had been keeping a male (in which owing to an injury to the hood the sperma-
tophore in the buccal recess protruded at the surface as in PI. LXXVII. fig. 5) under
careful observation for several days. Eventually the spermatophore was discharged from
its cyst, and I found it lying on the floor of the cage.
This is what inclines me to believe that it is necessary to work with a large
installation, a specially fitted-up aquarium of large size with concrete walls, in order to
rear the embryos of Nautilus successfully. I have no doubt that the shell and several
chambers are formed while the embryo is still in the egg-capsule, and that the young
Nautilus hatches out with the general facies of the adult.
I tried artificial fertilisation without success, and I do not think it is practicable.
The very young shell which is in all probability formed within the egg-capsule has been
called the nepionic shell, and may be easily recognised by its colour and finer grain in
young shells, being separated from the subsequent shell-substance by a well-marked suture,
which is visible in the umbilicus of the shells of N. macroinphalus and N. utnhilicatus.
The clear brown vitellus of the egg is surrounded by viscid colourless albumen
which accumulates at the poles. Between the yolk and the albumen there is a delicate
hyaline vitelline membrane. The vitellus is larger than that of any other known Cephalopod
When the females reach maturity, the bilabiate vagina and the nidamental gland
become enlarged, protuberant, and turgid, and the edge of the shell is thickened. The
nidamental gland frequently assumes a greenish colour. Thus a female N. jjompilius
with submature ovarian ova had a moderately developed, yellow-coloured nidamental gland
which had not become turgid. In another of the same age the yellow gland was tinged
with a greenish colour. In another the sub-turgid nidamental gland was of a dull
greenish or yellowish-gi'een colour. In a nearly mature female the nidamental gland
had a pronounced green colour. Apparently the gland does not always turn green, as
I have recorded instances of nearly mature females in which it exhibited a rich cream-
yellow coloration, the largest ovarian ova, in such specimens, attaining a length of
17"6 mm. and a maximum breadth of 15 mm.
The organ of Valenciennes also becomes turgid and functionally active with
abundant secretion of mucus at maturity.
The eggs of N. pompilius differ very slightly from those of N. macromphalus^ , and
• Willey, A., "The oviposition of Nautilus inacromphalus." P. R. Soc. London, 1897, Vol. 60, pp. 467—471, 6 figg.
810
VARIATION AND REGENERATION.
like the latter they are laid singly. The pectinate ridges are more prominent on the
outer capsule of the egg of N. pompilius, often having a frill-like appearance and pro-
jecting freely for a short distance at their anterior ends.
The capsules of the egg are often deficient in some respects, sometimes exposing
the vitellus through the gaping sutures of the
inner capsule (PI. LXXXIII. fig. 18), and in
the case of N. macroinphalus at Lifu I frequently
found fully formed capsules without any con-
tents, the latter not having escaped, but never
having been present. This also must I think
have been due to a derangement of the normal
reflexes, the result of capti\ity. The pectinate
ridges of the outer capsule of the egg seem
to correspond with the annulations of the infra-
buccal tentacles (PI. LXXIX. figg. 2 a and 6 a).
For further details concerning the egg-
capsules I may refer to the explanation of
figures 19—23 on Plate LXXXIII.
Fig. 14. Three eggs of N. macromphalus laid in
a row, with capsules coalcscent.
21. Vaei.\tion and Regexeratiox.
Although the old idea of the fixity of species has long been superseded, the fixity
of certain tj-pes seems to be a fact not open to question. Fixity of tv'pe is the ex-
pression of a completed evolution, and this is what is meant when a particular t\-pe
is said to be highly specialised in a given direction. Fixed t\-pes are verj' likely the
same as persistent types, and they have some properties in common amongst which are
their limited range of variation and their defective power of regeneration.
The variation in the number and disposition of the digital tentacles of Nautilus
is a negligible quantity, but the spadix exhibits ambidextral variation, being now on
the left side of the cephalopodium and now on the right. A variation of a similar
nature affects the origin of the main siphuncular arter}-, which sometimes arises as a
branch of the left division of the posterior pallial artery and sometimes from the right
division. The constitution of the spadix shows hardly any appreciable variation.
In one instance only did I meet with a situs inversus of the reproductive apparatus,
and this was the case with a male in which the vas deferens was developed on the
left side instead of on the right, wth the pyriform gland on the right instead of the
left side (PI. LXXVIII. fig. 2).
More than once or twice I found males in which the hood had been so bitten away
that the spermatophore lodged in the buccal recess was exposed to view. Sometimes
the front part of the hood is eaten away, at other times a piece is bitten out of
the middle of the hood just over the spermatophore (PI. LXXVII. figg. 5 and 6).
At first I attributed these injuries to the attacks of voracious fishes, but it is not
impossible that they are the results of nuptial combats. Without in any way relaxing
VARIATION AND REGENERATION.
811
its hold upon the shell, Nautilus can project its body from the shell with a darting
movement and protrude the buccal cone so as to seize hold of its food. I have seen
it seize a pra^\^l between its jaws in this way, and it would not be a difficult matter
for a female to bite into the hood of a male. However this may be, the point to
which I desire to draw attention is that the wounds heal up at the edges, but the
parts are not regenerated.
Table of Variations.
Sex
Spadix
Sipbuucular artery
Colour of Eeual ConcretiouB
1- i
2. „
3. „
4. „
5. „
6. „
7. „
8. „
9. „
10. „
Sinistral
Sinistral
Dextral
Sinistral
Sinistral
Sinistral
Sinistral
Sinistral
Sinistral
Sinistral
Sinistral
Sinistral
Dextral
Sinistral
Dextral
Dextral
Dextral
Sinistral
Dextral
Sinistral
11. 9
12. „
13. „
Sinistral
Dextral
Dextral
14. 6
15. „
Dextral
Dextral
Median
Dextral
16. <?
17. ?
18. S
19. „
20. „
Sinistral
Dextral
Sinistral
Sinistral
Dextral
Dextral
Dextral
Faint roseate (Rose-white)
Rose-white
Dark crimson
Wliite
Light scarlet
Amphioxus is another example of a relatively fixed type which presents little range
of variation, and less power of regeneration. Perhaps Chiton is another.
Sometimes the sheaths of the tentacles become perforated at some part of their
length, and the tentacle may then issue from the adventitious orifice instead of from
the apical orifice.
Not seldom among the shells of N. pompilius which are washed upon the reefs
of New Guinea specimens are found in which the umbilicus is incompletely closed'.
' I have figured some examples of this umbilical aberration in Quart. J. Micr. Set., sxxix., figures ou
PI. 13, 1896, pp. 227—230.
w. VI. 106
812
VARIATION AND REGENERATION.
Once a verj' abnormal shell was presented to me at Ralum in New Britain, in
which the shell was divided into two nearly equal halves by a submedian raphe, re-
minding one of the shell-slit of Pleurotomaria. The raphe extended back over the
Fig. 15. Malformed shell of ^V. pompilius.
involuted coils of the shell, and I am unable to decide whether it was due to an
injuiy to the mantle or to some congenital malformation.
WlLLiEY ZOOLOBICAL, ReSULTS,
Plate LXX
anterior
p,alUal artery-
Right
jballial vein
line of
insertioTh ,
of mantle.
outer
renal orqan
inner
pericardial
glandj .
outer
pericardial
jlarui.
Jiorn/y ___
girdle
Pallio- _.
vise, tig
sperm duct
(ves. sent.,
septal contoiuf:
igamentoiba bajici
from, funnel bwse.
, Columella, (shell/ J
muscle.
. ,- mantle edge .
Left
'pallial vein.
Inner
.- osphradium.
\
Y Inner renal
\ oraan
root of larger
bronchia/ .
outer
osjihradiAMn .
'-■root of
smaller brancTuo/ .
7jt ■- - shell muscle .
-' heart,
pynform body
testis
gijjard.
moescbne^
A"Wil'U3- W-ii na.t
West. Newman rlr
NAUTILUS MACR.OMPHAT t t •-;
WiLLEY Zoological Results.
Plate LXXVI.
p.Y.^
m.
,VCL.
-s.a.
6 dp.L
^
S.'K
CUC'
...gl.
.#■
/
'I
fi^..
\
'J'iBS'^"'
AWilleyScP.Highley del.
Wei.t.ls'ei-.iiiJJi ala-omc
HALIOTIS. HARPA. NAUTILUS.
DESCRIPTION OF PLATES LXXV— LXXXIII.
PLATE LXXV.
Nautilus macromphalus, $ . "View of pallio-visceral region from below with the organs
showing through by transparency.
Sketched from the fresh object, with natural colours, at Lifu, October 26th, 1896.
Each renal sac contains a bilobate renal follicle surrounded by the scarlet concretionary
substance. The liver has a dark purple colour.
PLATE LXXVI.
Fig. 1. If. pompilius, $ . Ventral view with reflected mantle, showing natural colours.
/.o. Funnel-organ.
a.i. Ala infundibuli.
X. Ligamentous limbus passing from funnel-organ over the surface of s.m. the shell-muscle.
m.e. Mantle-edge.
? Vulva.
hr. 1 and hr. 2. Branchiae.
osph. 1 and ospih. 2. The osphradia.
nid. Nidamental gland.
v.p.o. Viscero-pericardial orifice.
h. Bristle passed into the opening of the duct of the pyriform gland.
a. Anus.
ac. Accessory nidamental gland.
The renal orifices occur (when the mantle is reflected) in front of the bases of the
branchio-cardiac (eflerent branchial) vessels.
Fig. 2. Renal concretions ; the upper figure shows the concentric stratification under high
magnification.
Fig. 3. Harpa sp. from Lifu seen from below to show the divisions of the foot, the
semilunar propodium with sulcate border, the mesopodium and the metapodiuin. (Cf. text in
explanation.) Natural size.
FiGG. 4 and 5. Haliotis sp. from New Britain. Anterior end of animal with shell
removed, seen in ventral and dorsal view to illustrate the bilaminate, tentaculiferous epipodmm
(«p.) which is produced in front into a lobe wliich overlies the proboscis (buccal cone).
ni. Mouth.
/ Platypodium (foot).
e. Eye-stalk.
106—2
814 DESCRIPTION OF PLATES.
Between the bases of the cephalic tentacles occurs a free flap of the integument
resembling a nuchal membrane (Fig. 5). x 3.
N.B. — No portion of the mantle appears in either figure.
Fig. 6. iV. pomjnlius juv. Dorsal view of a portion of the mantle-region of the same
specimen as PI. LXXVII. fig. 4.
d.p.i. The line of insertion of the dorsal lobe of the mantle into the body-wall.
s.m. Shell-muscle.
an. Annulus.
s.a. Septal aponeurosis.
The succeeding figures, unless otherwise stated, relate indifferently to N. pompiliiis and
N. niacromphalus.
Fig. 7. Piece of the posterior region of the mantle viewed from the inner or coelomic
surface, to illustrate the connections of the siphuncle. The piece comprises the entire dorso-
ventral extent of the insertion of the genito intestinal ligament into the hinder wall of the
perivisceral coelom.
g.l. Genito-intestinal ligament (cut across).
p.a. Posterior pallial artery.
p.v. Posterior pallial vein.
s.a. Siphuncular artery (sinistral).
r.s. Radical sinus of the siphuncle (pallio-siphonal sinus).
s. Siphuncle.
Fig. 8. Similar piece from another specimen. The posterior pallial vein and pallio-siphonal
sinus have been laid open and a bristle passed into the siphuncular vein (axial cavity of
siphuncle). The siphuncular artery (dextral) is seen in the floor of the pallio-siphonal sinus.
Fig. 9. Portion of a bisected shell of T. pompilius to illustrate the relation between
the growth-lines of the shell (shell-lines), the concentric muscle-lines and the posterior border
of the actual muscle-scar and annulus. An asterisk is placed opposite to the linea limitans
of the muscular impression. The last-formed septum is thin and fragile, thickening not having
yet taken place.
PLATE LXXVII.
Fig. 1. Frontal view of the fundus of the animal-chamber of a shell of N. pompilius.
1. Annular impression.
2. Septal suture or impression of the septal aponeurosis of the mantle.
3. Muscle-scar.
4. Border of the nacreous layer deposited by the dorsal lobe of the mantle over the
black varnish which covers the normally banded periostracum of the anfractus of the shell.
Fig. 2. Specimen of female N. macromphalus with spermatophore {sp.) protruding from
the oral sheath.
Fig. 3. N. macromphalus adhering to the substratum in a vertical position by means
of its tentacles. More usually its position, when attached to any surface, is more or less
horizontal, with the adhesive tentacles spreading about in all directions.
WiLiiEY Zoological Results.
Plate LXXVII
"^
?-' ~^^
t
i-r
M
rn
ffl^,f}errT^\^
.M" .r
t^
y
%
&.
\
'few
P.HigHey&AiVjiley dffi
NAUTILUS.
West.Newmaa litli
WlLLEY ZoOLiOGICAL RESULTS.
Plate LXX.VI11
^
H' h
»*>"
/'
,/
J
Edvnn Wilson del ,
West,Newina3i lith
NAUTILUS
DESCRIPTION OF PLATES. 815
a.i. Ala itifundibuli in diastole.
n.m. Nuchal membrane detached from the anfractus of the shell.
e. Eye with the ophthalmic tentacles on either side of it.
Fig. 4. Dorsal view of young N. pompilius to show relations of hood, nuchal membrane
and mantle in the nuchal region.
7i.m. Nuchal membrane passing round on either side to coalesce with the alae in/undihuli.
b.w. Portion of body-wall between the base of the nuchal membrane and the line of
insertion of the dorsal lobe of the mantle.
d.p.f. Dorsal lobe of mantle turned back.
s.m. Shell-muscle.
si. Siphuncle.
Fig. .5. Dorsal view of cephalic region of a male specimen of JV. rnacromphalus in which
the right anterior half of the hood had been apparently bitten away and the edges healed up.
/. Funnel.
in. Buccal cone.
sp. Cyst of spermatophore, with the apices of some of the extrabuccal tentacles exposed
on either side.
s. Spadix.
Fig. 6. Similar view of another specimen in which a piece had been bitten out of the
hood so as to expose the entire cyst of the spermatophore (sp.), which is held by the two
dorsal tentacles of the extrabuccal series on each side.
PLATE LXXVIII.
FiGG. 1 and 2. Two male lY. pompilius from below with mantle drawn back, showing
(in Fig. 2) situs inversus of the genital apparatus. These specimens are now in the Zoological
Museum of the University of Cambridge.
v.n. Visceral nerves.
OS. Submedian osphradia, separated by an interval in Fig. 1, coalescent in Fig. 2.
n.s. Needhamian sac of the vas deferens.
Fig. 3. Side view of cephalopodial region of N. uiahilicatus to show the character of
the hood.
a. Sulcate tentacle of spadix (10th tentacle of the extrabuccal .series).
b. Tessellate tentacle of spadix (11th of the .series).
p.g. Preputial gland.
e. Root of eye, wliich had been removed by means unknown to me ; a bristle is passed
into the olfactory labyrinth.
/. Funnel
a.i. Ala infuiidibuli.
The digital tentacles of the left side have been cut away in order to expose the spadix.
Fig. 4. Cephalopodium of ^V. macmmphalus with erected spadix. The hood has been
divided in the middle line. Specimen deposited in the Museum at Cambridge.
g. Glans of spadix (9th tentacle of extrabuccal series).
sp. Cyst containing spermatophore.
c.e. Cut edge of hood,
e. Eye.
816 DESCRIPTIOX OF PLATES.
PLATE LXXIX.
Fig. 1. Portion of shell of ^"'. niacromphalus showing encrusting organisms in and near
the umbilicus.
Fig. 2. Egg of XautUus niacromphalus, showing fenestrations and pectinations of the
outer capsule. There is a deep median notch extending from the front end through about
half the length of the capsule, x li.
Fig. 2 a. Portion of one of the pectinate ridges of preceding. These ridges seem to
correspond with the annulations of the iuf rabuccal tentacles (cf. tig. 6 a).
Fig. 3. Same egg seen from the lower surface, showing the area of fixation.
Fig. 4. Secretory products of the nidamental gland. The oval bodies are formed in
large bladder-like cysts. Zeiss 3 D, Cam. luc. ..Drawn from fresh object at Lifu, 9/2/97.
Fig. 5. Infrabuccal apparatus of immature female {N. pompilius).
Fig. 6. Infrabuccal apparatus of mature female (X. pompilius) with the subjacent portion
of the cephalopodium.
0. Organ of Owen.
V. Organ of Valenciennes.
19. Nineteenth digital tentacle of the outer whorl.
Fig. 6 a. Enlarged drawing of one of the infrabuccal tentacles of preceding.
Fig. 6 b. Apex of one of the infrabuccal tentacles of same.
Fig. 7. Anterior portion of cephalopodium (X. pompilius ? ) with hood partially bisected
and the flaps turned aside to show the inner whorl of tentacles embracing the buccal cone.
/. Funnel.
0. Organ of Valenciennes.
i.a. Infrabuccal apparatus.
e.l. Extrabuccal lobe.
Fig. 8. Buccal cone of female in side view, -n-ith buccal membrane pressed back to expose
the jaws.
i.m. Lower jaw.
r. Teeth of radula.
s.m. Upper jaw.
i.a. Infrabuccal apparatus.
p. Marginal papillae of buccal membrane.
Fig. 9. Spadix of X. niacromphalus (same as in PI. LXXVIII. fig. 4) in frontal view.
ff. Glans.
a. Sulcate tentacle.
b. Tessellate tentacle.
jj.g. Preputial gland.
Fig. 9 a. Portion of sulcate tentacle enlarged, showing glandular pits.
Fig. 9 b. Similar portion of tesseUate tentacle, showing glandular crypts.
Fig. 10. Same spadix from left side. The maximum length of this spadix is 65 mm.;
the glans projects 28'.3 mm. beyond its sheath ; maximum width of glans, 25 mm.
WltLEY ZoOLOGICAl. ReSULTS.
Plate LXXIX.
9'
West Jlewmaa litli
NAUTILUS.
WiLLEY Zoological "Results.
Tlate LXXX.
P Highley dtl et-lilh ,fcW,lley Je. .
ti. 21
WILLEY NAUTILUS .
HanKart imp.
DESCRIPTION OF PLATES. 817
Fig. 11. Infrabuccal apparatus of female (iV". viacromphalus) in state of erection. It
seems likely that the tentacles of this organ take an active part in fashioning the outer
capsule of the egg. (Cf. figg. 2 a and 6« of this plate.)
Fig. 12. Infrabuccal organ (organ of Van der Hoeven) of the youngest male {N. pompilius)
obtained, x 20.
Fig. 13. One of the outermost digital tentacles, to show the protuberant ridges towards
the apex, x 2.
Fig. 14. Portion of an ophthalmic tentacle showing ciliated lamellae. (Reduced from
figure published in Quart. J. Micr. Sci. 1897.) . .
PLATE LXXX. {N. pompilius.)
Figg. 1 and 2. Views of the right (1) and left (2) extrabuccal series of tentacles in
the youngest male obtained. The ventral group of four tentacles represents the spadix or
the anti-spadix, but is, at this stage, exactly equal on both sides of the cephalopodium.
«. Cut edge of ventral symphysis.
Fig. 3. Young (sinistral) spadix showing unmodified apices of the tentacles protruding
from their sheaths, x 3.
[The middle length of the hood of the original specimen was about 33 mm., and width,
measured over the eyes, about the same.]
The letter g is placed upon the sheath of the first tentacle or ijlans of the spadix.
i. Cut edge of intercoronal membrane which was removed in order to expose the entire
spadi.x which was sunk deeply behind it.
s. Cut edge of ventral symphysis.
Fig. 4. Outer aspect of same, showing f. the free tentacle of the spadix, partially
concealed behind the membranous e.xpansion of the sheath of the glans.
Fig. 5. The same, with sheaths of tlie tentacles laid open and the free tentacle {/.) cut off.
g. Glans.
Fig. 6. Another (dextral) spadix rather older than preceding, showing commencing
reduction of the apical portion of the glans (g.).
f. Free tentacle of spadix.
Figg. 7 and 8. Spadix (7) and anti-spadix (8) of an older specimen (middle length of
hood nearly 40 mm., width across the eyes about 38 mm.), x 3.
The free tentacle of the spadix is, at this stage, the most prominent member of the series.
i. Cut edge of intercoronal membrane.
s. Cut edge of ventral symphysis, with the nineteenth digital tentacle of the outer
whorl of the cephalopodium, beside it.
Fig. 9. Inner surface of the ventral symphysis of a young female (measuring 66 mm.
from the root of the siphuncle to the anterior border of the hood), showing the commencing
formation of the lamellae of the organ of Valenciennes. Natural size.
Fig. 10. The right extrabuccal lobe of the same specimen; free dorsal border on tlie
left, cut edge below and on tlie right of the figure. The tentacles have attained their normal
development (compare figg. 1 and 2). x 2.
818 DESCRIPTION OF PLATES.
Fig. 11. Infrabuccal apparatus of the same, presenting a prominent, transversely extended
"organ of Owen" and lateral tentacles in a rudimentary condition, x i.
Fig. 12. Portion of the inner surface of the ventral pallial fold of same, to show the
primordia of the sheath of the nidamental gland ; the lamellae of the gland (compare
PL LXXVI. fig. 1) have not yet commenced to form, x 2.
o. Submedian osphradium.
Fig. 13. Infrabuccal organ of adult male from the ventral (aboral) aspect with the
membranous sheath partially reflected to expose the lamellae (compare PI. LXXIX. fig. 12).
The total length of the organ was "26 mm. and the width 14 mm.
Fig. 14.. Similar infrabuccal organ from another specimen viewed from the dorsal (adoral)
aspect, with the wall cut away so as to expose the cavity of the gland and the glandular
pits on the surface of the lameUigerous lobes (l.).
h. Cut edge of the fleshy wall of the gland ; this wall is derived from, and adherent
to the periphery of the lameUigerous lobes and peduncle of the organ, leaving only a narrow
median passage (on the dorsal side) from the cavity of the gland to the exterior. In the
dissection here figured the dorsal portion of the membranous sheath of the organ was divided
by an incision passing through the median (dorsal) orifice.
Fig. 15. The same bisected so as to expose the inner or mesial surface of the left
half of the gland, showing the free inner borders of the lamellae and of the lameUigerous
lobe (/.).
p. Cut surface of the median portion or peduncle of the organ.
m} and rrv. Cut edges of the dorsal and ventral portions of the membranous sheath of
the organ ; the sheath is a derivative of the buccal membrane, i.e. the integument of the buccal
cone.
Fig. 16. Male genital duct removed from the body.
6. Ostium abdominale.
gl. Glandular portion (phorogenic gland, containing a labyrinthine lumen in which the
spermatophore is formed) of tlie vesicula semimdis, usually called the accessory gland.
a. Thin-walled antrum ; in front of this region, the duct passes through the integument
by a narrow neck and then dilates to form n, the Needhamian vesicle, which is connected
with }), the penis or tenninal portion of the duct, by a narrow sphincter.
Total length of vesicula sf.mitudis (gland + antrum) 26 mm.
Fig. 17. Portion of the same with the antrum laid open showing the orifice perforating
its floor by which the phorogenic gland communicates with it ; a portion of a spermatophore
is projecting freely through the orifice into the antrum. The roof of the antrum is perforated
by another minute orifice with somewhat tumid lips, leading into the Needhamian vesicle.
Letters as in preceding figure.
Fig. is. Another' portion of the same. A piece of the wall of the Needhamian vesicle
has been removed in order to expose the spermatophore which it contains and the saddle-
shaped septum upon which the spermatophore rests.
Letters as in fig. 16.
Fig. 19. The spermatophore removed from the Needhamian vesicle, showing the notch
occasioned by the septum of the vesicle.
Fig. 20. Pallio-visceral region of youngest male from below, chiefly to illustrate the
DESCRIPTION OF PLATES. 819
genital or, as called in the text, the geuito-intestinal ligament, which rejjresents a median
mesentery, x 3.
t. Testis, derived from and adherent to the right side of the mesentery , on the opposite
side is the gastric ligament passing across to be inserted into the wall of the stomach {st.).
p.a. Posterior pallial artery, seen traversing the middle pallio- visceral foutanelle (cf.
PI. LXXXIl. fig. 7).
SI. Siphuncle.
FiGG. 21 and 22, Details of preceding, to elucidate the relations of the transverse and
longitudinal loops of the intestine. Fig. 21 shows a portion of the intestine and right liver
in normal juxtaposition. In fig. 22, the liver has been cut away so as to expose the
intestine and the longitudinal loop of the latter has been turned forwards.
t.i. Transverse loop of the intestine.
d.i. and a.i. Descending and ascending limbs of the intestine.
oe. Narrow portion of the oesophagus proceeding from the fundus of the crop to the
stomach (St.).
py. Pyloric caecum with bile-duct opening into its apex.
h.d. Bile-duct, above or behind which the intestine passes.
Fig. 23. Pallio-visceral region of another specimen older than that of fig. 20, but showing
the same primary relations of the mesentery.
y. Pyriform gland.
St. Stomach traversed in this specimen by constrictions.
l).a. Posterior pallial artery.
d.i. and a.i. Intestine.
r.l. Right liver.
PLATE LXXXI.
Fig. 1. N. pompilius. Dissection of a relaxed specimen from the right side, showing
the relation of the capito-pedal cartilage to the siphonopodium and to the cephalopodium.
The alcie infundihuU have been cut away.
/. Funnel {crura infundibuli).
I. Lhnbus.
c. Cartilage.
V. Infundibular vein.
a. Infundibular artery.
r. Rhinophore.
o.a. and o.p. Anterior and posterior ophthalmic tentacles.
e. Eye.
1 — 11. sheaths of digital tentacles; that of the second tentacle (2) comprises an anterior
portion forming part of the hood and a posterior portion ; the third sheath (3) is wedged
in between 2 and 4, only its anterior end being visible in this view.
Fig. 2. Diagram of Naidilus to illustrate the theory of orientation. The dotted line
denote.s the presumptive antero-posterior axis.
I. Limhus.
a.i. Ala infwidibuli.
s.in. Shell-muscle.
m. Mantle.
s.a. Septal aponeurosis.
w. VI. 107
820 DESCRIPTION OF PLATES.
FiGG. 3 and i. Diagrammatic sections through the cephalopodium of female (3) and
male (4) to elucidate the tentacular formula suggested in the text. The ophthalmic tentacles
are not included in the diagrams. On the left of fig. 4 there is the scheme of enumeration
of the digital tentacles of the outer whorl, the tentacles being here regarded as marginal
appendages of a sinuous pleural fold or epipodium.
c.p. Marginal papillae of the mouth.
m.i. Inferior mandible.
s.s. Superior symphysis of the cephalopodium.
i.s. Inferior symphysis.
/. Funnel.
ext.t. Outer whorl of the cephalopodium.
mt.L + inf.o. Inner whorl.
in/.o. Infrabuccal organ.
Fig. 5. ^\' pompilms ^. Dissection from below to illustrate, among other points, the
ventral symphysis of the cephalopodium which presents a deep concavity, f.i. the fossa
infundibularis, to receive the funnel. The numbers are placed upon the slieaths of the
tentacles involved in this view, in accordance with the scheme of enumeration here adopted.
The funnel has been sliced away to its roots and the vena cava and endochondral sinus
laid open to expose the fenestrations in their dorsal wall.
r.i. Radix infundibuli.
c.i. Cartilage.
a.i. Ala infundibuli.
v.i. Vena cava.
s.m. Shell-muscle (cut across).
n.s. Needhamian vesicle.
e. Eye.
Fig. 6. N. pompilius. Dissection of the cephalic sensory complex from the right side,
showing also the origin in sequence from the coronal (pleuro-pedal) ganglion, of the nerves
of the sixth, seventh and eighth digital tentacles.
a. Infundibular artery traversing the cartilage.
a.i. Cut surface of ala infundibuli.
c. Cartilage.
c.v. Cerebro- visceral nerve-centre.
h. Cut surface of hood.
l. Limbus infundibrdi.
n. Olfactory labyrinth (rhinophore) laid open.
71.711. Nuchal membrane.
71.71. Olfactory nerve.
o.p. Posterior ophthalmic tentacle.
op.n. Optic nerves.
ot. Otocyst, lying over against the coronal ganglion, concealing the mass of the latter
(cf. PI. LXXXII. fig. G).
s.m. Surface of shell-muscle.
('. Infundibular vein.
Fig. 7. S. pompilius. Dissection of the nuchal complex from above. The hood and
nuchal membrane have been divided along the mid-dorsal line and the halves drawn apart.
WiLLEY Zoological Results .
Plate LXXXl
P.HigWey lifli . Wdlcy del .
nfiTiacii-l, imp.
WILLEY, NAUTILUS
DESCRIPTION OF PLATES. 821
exposing the peristomial and peri-oesophageal haemocoels and the diaphragm which separates
them from one another behind the cerebral capsule.
a.i. Dorso-posterior portion of ala injundibuli.
ao.s. Aorta superior.
a.p.a. Anterior proventricular artery.
b.c. Buccal cone.
c.c. Cerebral capsule attached to neighbouring structures by conjunctive trabeculae and
produced in front as a broad lobe.
cr. Crop.
m.i. Inferior mandible.
n.m. Nuchal membrane.
pr.m. Protractor muscle of the buccal cone.
r.i.m. and r.s.m. The dorsal retractor muscles of the buccal cone, r.i.ni. attached to
the inferior, r.s.m. to the superior mandible.
N.B. The cerebral commissure only occupies the posterior band-like portion of the massive
fibrous capsule. It is possible that this fibrous capsule represents a primordial condition of
the cartilaginous capsule of Dibranchs which is not, I think, homologous with the capito-
pedal cartilage of J\'autihts, the latter being essentially a funnel (siphonal) cartilage and not
a cranial cartilage.
PLATE LXXXII.
Fig. 1. iV. rnacromphalus S juv. Innervation of the infrabuccal organ (organ of V^an der
Hoeven). The nerve to each laraelligerous lobe (i.e. to each half of the organ) ends abruptly,
giving off the lamellar nerves as shown on the right side only (left of the figure).
m. Shallow secondary fold of the buccal membrane which would form the adoral portion
of the sheath of the organ.
The middle length of the hood of the specimen was 52 mm. ; the longer diameter of
each lobe of the infrabuccal organ measured 5 mm.
Fig. 2. A'', pompilius <$ juv. Innervation of infrabuccal organ ascertained from an injected
specimen.
a, b and c. Branches of the infrabuccal artery which accompanies the nerve.
Middle length of hood 65 -5 mm. ; length of infrabuccal organ 6 mm.
Fig. 3. Topography of the infrabuccal organ of the male. The fore-part of the funnel
has been cut away and the ventral symphysis di\'ided down the middle, thus exposing the
ventral portion of the peristomial haemocoel, the organs which project into it and the nerves
and vessels which traverse it. The dotted line represents the limit of the fundus of the
external fossa buccalis
p.c. Capsule of the coronal (pleuro-pedal) commissure, from which a small median artery
issues and, passing freely across the haemocoel, enters the fundus of the infrabuccal organ.
n. Infrabuccal nerve accompanied by the artery.
l.i. M. levator infundlbuli.
c.i. Cartilage.
a.i. Ala infundibuli.
Fig. 4. Somewhat similar dissection of female {N. macromphalus). The fundus of the
peristomial haemocoel is exposed from the ventral aspect.
107—2
822 DESCRIPTION OF PLATES.
i.o. Infrabuccal organ (only the sheaths of the tentacles indicated on either side of the
organ of Owen).
e.b. Extrabuccal lobe.
c.e. Cut edge of buccal membrane.
i.n. Infrabuccal nerve.
n./. Infundibular nerve.
p.c. Coronal (pleuro-pedal) commissure.
a.c. Arcus cartilaginis.
i.r. Ventral retractor muscle.
N.B. The actual distance between the roots of the infundibular nerves was 14 mm. ;
between the infundibular and infrabuccal nerves on each side, 4 mm.
Fig. 5. Another dissection (If. pompilius <S) exposing the fundus of the peristomial
haemocoel ; the funnel and oval sheath (cephalopodium) have been sliced away and the buccal
cone turned back.
Letters, commencing from left side of figure : —
r.a. Recurrent tentacular artery.
o.s. Orifice of ophthalmic sinus.
c.s. Orifice of spadical sinus (a spacious reservoir surrounding the base of the spadix,
probably assisting, by blood-pressure, in the protrusion of the latter).
i.n. Infrabuccal nerve.
n.i. Infundibular nerve.
a.i. Arcus cartilaginis.
c.c. Coronal commissure.
i.r. Ventral retractor.
c.i. Cartilage.
l.i. Levator infundihuli.
e. Eye.
h. Hood.
Between a.c. and c.c. lies the passage leading from the peristomial haemocoel into the
endochondral sinus.
Fig. 6. Dissection resembling that of PI. LXXXI. fig. 6, but carried further. The
ophthalmic sinus is cut open, the rhinophore pulled aside, and the otocyst turned back.
a.o.t. Anterior ophthalmic tentacle.
a.o.n. Anterior ophthalmic nerve.
I. Hood-nerve.
II. — VIII. Digital nerves. (Between II. and V. occurs the common root of III. and IV.)
V. Visceral nerve-collar cut across.
0. Otocyst, turned back to expose the massive coronal ganglion.
o.g. Optic nerve-ganglion complex.
p.o.n. Postei'ior ophthalmic nerve, taking a recurrent course in the wall of the ophthalmic
sinus, mediad of the optic ganglion (the posterior ophthalmic tentacle itself is quite concealed
behind the eye in this view except for the apex which happens to project forwards between
the eye and the hood).
r. Rhinophore.
e. Eye.
Wllley. Zoological Insult,
P HiaK'lsvae'i (i Ktii .ViMlev «!
Han'harC imp
WILLEY. NAUTILUS .
DESCRIPTION OP PLATES. 823
Fig. 7. Dissection of the pericardium from below to show the three fontanelles in the
pallio-visceral ligament leading from the pericardium to the perivisceral coelom. [Reduced
from Quart. J. Micr. Sc, Vol. XL. 1897, PL II. fig. 2.]
p.l. Pericardial ligaments.
p.g. Pericardial glands.
b.c. Branchio-cardiac vessels.
a.p.a. Anterior pallial artery.
py. Pyriform gland.
g.d. Genital duct.
r.s. Outer renal sac (cf. PI. LXXV.).
The pyriform body and its ligament form part of the outer border of the middle
fontanelle and here somewhat encroach upon it so as to disgui.se its full extent. The
posterior pallial artery is seen crossing the ventral surface of the heart, disappearing behind
the pyriform ligament through the middle fontanelle. The heart is liound to the pallio-
visceral ligament (which constitutes the fundus of the pericardium) by the cardiac ligament.
Fig. S. Dissection of the branchial apparatus of the right side. The membranous portion
of the mantle has been turned back and partly removed. The efferent trunk (b.c.') of the
larger gill has been cut across and the gill itself turned partly round to present the atferent
surface which has been laid open to expose the cavity of the reno-branchial vessel {r.b.~).
The portion of the latter which lies within the pericardium is left intact.
c.e. Cut edge of the mantle at the junction of its membranous and visceral portions.
r.s. and p.g. Renal sac and pericardial gland of the macrobranchiate segment.
b.c.'^ Extra-pericardial and intra-pericardial portions of the branchio-cardiac trunk.
b.c' Branchio-cardiac vessel of the lesser gill (seen lying across in front of the greater gill).
V. Heart.
p.v.l. Pallio-visceral ligament.
p.l. Pericardial ligaments.
a. Position of annulus.
s. Position of septal aponeurosis.
ni. Mantle.
Fig. 9. Portion of body behind the funnel to show veins entering the vena cava.
f.o. Funnel-organ.
I. Liinbus or ligament passing from f.o. over the surface of the shell-muscles.
Fig. 10. Ventral view of the pallio-visceral region of N. pompilius, to show the paired
pallial veins as displayed by automatic injection with air or gas after removal of the animal
from the shell. The finest ramifications of the veins throughout the substance of the
membranous portion of the mantle are thus revealed, but only the main branches are shown
in the drawing.
PLATE LXXXIII.
FiGG. 1 and 1 a. Amoeljocytes _ (blood-corpuscles) of N. macrnmphalus. They undergo
relatively rapid amoeboid changes of form ; some contain fine granules, but most of them are
laden with coarse, bluish, refringent granules which dissolve away in acetic acid leaving a
spongy plasmatic reticulum. The cells vary in size and tend to accumulate in masses on
the slide, sometimes appearing to form plasmodia (tig. 1 a). When killed and stained with
824 DESCRIPTION OF PLATES.
acetic acid and alum cochineal the nucleus presents a chromatic reticulum. Zeiss Oc. 3, Obj. J,
water imm., Cam. luc.
Fig. 2. Otocones. Zeiss 3 J, Cam. luc.
Fig. 3. Sagittal section (parallel to long axis of body) through one of the submedian
osphradia. Only the epidermis and nerves are drawn. The limits of the sensory epithelium
are indicated by slight elevations near the borders of the figure. A large nerve proceeding
through the conjunctive framework from below is seen to branch and then penetrate through
the basement-membrane into the substance of the epidermis, where it terminates in an intra-
epidermal fibrillar plexus. Zeiss 3 A.
b.in. Basement-membrane.
n. Nerves with subcutaneous and intra-epidermal course.
Fig. 4. Portion of a similar section more highly magnified, showing a remarkable intra-
epidermal fibrillar plexus. Zeiss 3 D, Cam. luc.
Fig. 5. Tangential section through the epidermis of a submedian osphradium taken
through one of tlie angles made by the papilliform lobes (which arch over and protect the
sensory surface of each submedian osphradium) with the base of the osphradial groove. The
ramifying intra-epidermal fibrillar plexus is well shown, and there are also indications of the
occurrence of peripheral ganglion-cells. Zeiss 3 C, Cam. luc.
Fig. 6. Papilla of an interbranchial osphradium showing cilia. Drawn from the living
object and reproduced from Quart. J. Micr. Sc, 1897.
Fig. 7. Section through an interbranchial osphradium cutting the papilla lengthwise,
i.e. at right angles to the long axis of the body, showing the papilla itself and the subjacent
surface of the mantle. The sensory surface occupies a well-defined tract about the angle
made by the papilla with the mantle. Cilia not shown in the drawing.
o.p. Apex of the osphradial papilla.
n. Subcutaneous and intra-epidermal nerves.
Fig. 8. Enlarged view of the intra-epidermal termination of a nerve (n.) in an inter-
branchial osphradium.
Fig. 9. Transverse section through the siphuncle showing the central sinus (siphuncular
vein) and peripheral lacunae communicating with it.
a. Main artery of the siphuncle.
d. and v. Dorsal and ventral surfaces of the siphuncle.
Fig. 10. Portion of the wall of the siphuncle showing the intra-epidermal prolongations
of the subcutaneous lacunae. Zeiss 2 D, Cam. luc.
s. Subcutaneous blood-spaces or lacunae.
t. Trabeculae.
Fig. II. Longitudinal section through siphuncle, showing the axial vein oommunicatLng
with the peripheral lacunae.
«. Axial sinus or vein ; (the index lines pointing to the sides of the sinus should not
have been inserted).
Fig. 12. Portion of wall of preceding, showing the longitudinal course of one of the
intra-epidermal blood-channels.
s. Venous space.
WiLLEY Zoological Eesults .
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Plate LX)0(I11.
Han hart imp
us
DESCRIPTION OF PLATES. 825
Fig. 13. Bete mirabile seen through the epidermis of the funnel-organ in the fresh state.
Fig. 14. Portion of endochondral sinus showing a valve arching over the capito-pedal
orifice. An arrow is passed from the tubular sinus in which tlie levator muscle of the funnel
lies, into the endochondral sinus.
c. Cartilage.
V. Valve.
Fig. 1.5. Ovarian ovum of an injected specimen, removed from the body and \iewed
from the animal pole showing the raniitications of the genital artery in the follicle.
Fig. 16. Enlarged view of portion of preceding.
Fig. 17. Section through an ovarian ovum, showing the intra-lecithal processes of the
follicle.
Fig. 18. Abnormal egg of N. inacromphalus, with defective capsule exposing the viteUus
at three places. The outer capsule was only present on one side of the egg, namely, on
the side opposite to that figured.
Fig. 19. Inner capsule of an egg of ..\'. pomjiiliun from below, showing clear mesial
longitudinal tract separating the striations of the remaining surface. The outer capsule has
been removed except in the region of its median and lateral attachments to the inner capsule.
Fig. 20. Pear-shaped inner capsule of egg of If. macromphalus from below. The outer
capsule has been removed except at the hinder end in the region of the irregular disc of
attachment which presents a spongy cavernous surface. The striations of the inner capsule
tend to run longitudinally below and transversely on the upper side of the egg.
Fig. 21. Egg of N. macromphalus from above, with the longitudinal slit in the upper
wall of the outer capsule widened out so as to expose the inner capsule with its striated
surface. Reproduced from P. R. Soc. London, Vol. lx. 1897, p. 470, tig. 4.
Fig. 22. N. macromphalus. Inner capsule of egg seen from below (i.e. from the side
directed towards the disc of attachment). Half the lower wall of the capsule has been
removed by slitting along one of the lateral sutures and along the median groove (depressed
longitudinal tract) to show the brown-coloured vitellus lying in the capsule. The lateral
sutures are continued into each other at the front of the capsule so that the inner capsule
is really bivalvate. The germinal disc (not indicated) occurs at the anterior pole of the
vitellus, the orientation of the laid egg coinciding with tliat of the ovarian egg (cf. text-
fig. 13).
Fig. 23. N. macromphalus. Inner capsule of egg in side view, showing tlie dorsal
ridge with its anterior terminal process and the lateral suture. A. fragment of tiie outer
capsule remains at the hinder end, the rest having been cut away.
Fig. 24. Enlarged view of one of the nascent tentacles of the extrabuccal series shown
in PI. LXXX. tigg. 1 and 2.
The tentacle appears as a minute papilla and the sheath seems to grow round it as a
dermal fold with a simple border in fr<mt and a deep notch (po.ssibly indicating concrescence
of the two halves of the fold) at the back of the tentacle. Zeiss 3 A.
Fig. 25. Macroscopic view of the eye in vertical section. The piano of section lies
on one side of the pinhole aperture so that this is seen intact from tlie interior of the
optic cup. The bacillary layer representing the vitreous body forms a thin sheet over tlie
826 DESCRIPTION OF PLATES,
black retina except over the inner surface of the opaque cornea which appears as a black
disc perforated by the pinhole orifice.
The dorsal border of the eye is smooth and convex, the ventral border limbate. x 3.
r. Rhinophore.
Fig. 26. View of the anterior portion of the peri-oesophageal haemocoel. The body has
been divided in the nuchal region. The visceral nerves are seen radiating out from the
visceral ganglionic commissure.
d. Diaphragm (the darkly shaded area in front of and above the bifurcating aorta)
separating the peristomial haemocoel from the peri-oesophageal haemocoel.
a. Aorta.
n.in. Nuchal membrane.
h. Hood.
oe. Oesophagus.
8.m. Shell-muscle.
v.c. Vena cava.
f.o. Funnel-organ.
a.i. Ala infancUbuli.
Fig. 27. The funnel with the flaps pressed open to expose its internal surface. The
regions of the crura and alae are separated by the anterior borders of the cartilage appearing
as oblique, wliitish bauds. Between the crura occurs the tongue-shaped valve, the free border
of whicli is approximately concentric with the anterior border of the funnel.
f.o. Funnel-organ.
Fig. 28. Dissection of an injected specimen to exhibit the recurrent course of the
tentacular artery. (Cf. also PI. LXXXII., fig. 5.)
a. Aorta.
r.m. Roots of dorsal retractor muscles.
c.e. Cerebral ganglionic commissure cut across ; the root.s of the two stomatogastric
nerves of the right side are seen beyond the cut edge, but the demarcation of the cerebral
commissure could not be well indicated in this drawing.
g. Coronal (pleuro-pedal) ganglion.
r.a. Recurrent tentacular artery.
Fig. 29. Dissection of the buccal cone from the right side, to illustrate the topography
of the stomatogastric system.
r.m. Retractor muscle.
«. Buccal artery.
r. Radular caecum.
n'. and n". Stomatogastric nerves meeting in the pharyngeal ganglion below the border
of the wing-like expansion of the lower mandible {ir. is cut across shortly before the union).
h.c.^ and b.cr Buccal commissures proceeding from the pharyngeal ganglion.
INDEX.
n
Abdomen, 798 (see also Vi>eeral sac)
Abdominal pores, 805
Accessory -gland ( <? ), 7S7
Accessory sexual organs, 776, 777
Aoephala, 788, 790
Acetabula, 768
Acetabular Theory, 800
Adams, A., 809
Afferent branchial vessels (see Beno-
brauchial vessels)
Affinities, 805
Air-chambers, 747, 798
Ala,' iiifundibiUi, 753, 765, 772,
783, 790, 802
Albumen, 809
Ammonites, 737, 743
Amoebocytes, 783, 789
Amphineura, 797, 805
Anastomosis, 763
Anfractus of shell, 740, 748, 751,767
Annular lobe, 767, 771, 776
Annulus, 747, 749
Antero-posterior axis, 797, 805
Anti-spadix, 776
Antrum (i ), 787
Anus, 753, 768
Aorta inferior, 755
Aorta superior (see Systemic aorta)
Aplaeophora, 806
Aponeurosis, 749, 750
Appellor, A., 748, 749, 750, 751,
760, 807
Area noae, 792
Arcris cartilaginis, 766, 783
Argonauta, 808
Arms of Cephalopoda, 768, 798,
800, 802, 803
Atlanta, 795
Autotomy (metapodium of Harpa),
795
Axial nerve, 768
Axis, 764, 797, 806
W. VI.
Bait, 774
Bather, F. A.. 742, 743, 748
Belemnites, 737
Bennett, G., 737, 743
Bles, E. J., 805
Blood, 783
Boulenger, G. A., 797
Bourne, A. G., 737, 740, 754, 767,
791
Boutan, L., 796
Branchiae, 753, 788, 804
Branchial folia, 789
Branchial gland, 789
Branchial skeleton. 789
Branchial Theory, 800
Branchio-cardiac vessels, 7.55, 781,
788, 804
Branchio-o»phradial arteries, 756
Brock, J., 795, 807
Brooks, W. K., 788, 802, 803
Buccal cone, 765, 766, 767, 781
Buckman, S. S.. 743
Callus of shell, 743, 753
Capito-pedal cartilage, 764, 766,
783, 797, 803
Capito-pedal (venous) orifices and
valves, 783
Cardiac ligament, 781
Carina eartilaginis, 7()5
Cartilage, 763
CephaUc arteries, 782
Cephalic cartilage, 764, 766
Cephalic sense-organs, 793, 803
Cephalic Theory, 801
Cephalisation, 806
Cephalopoda, 740, 765, 766, 790,
796, 799, 805
Cephalopodium, 763, 765, 767, 769,
793, 796, 797, 802, 803, 806
Cephalothorax, S06
Cerebral nerves, 774
Cerebro-visceral system, 781, 791,
804
Chiton, 797, 799, 806
Cilia, 773, 788, 791, 802
Circulus pallialis, 763
Cirri of Cirroteuthis, 800
Cirroteuthis, 800
Coelom, 758, 805
Columellar artery, 782
Columellar muscles (see SheU-
muscles)
Compressor muscles, 765
Conchy olin, 749
Concrescence, 771, 803
Cooke, A. H., 795
Corium, 739
Coronal ganglion, 775, 783, 794.
804
Costate tentacle (of spadix), 777
Crick, G. C, 743, 746
Crura infundibuli, 764, 765, 802
Ctenidium, 789
Cuvier, Baron, 737, 738
Dean, B., 738, 740
Dentalium, 803
Development (of accessory sexual
organs), 777, 778
Diaphragm (separating the peri-
stomial haemocoel from the
peri-oesophageal haemocoel, 780,
782
Dibranchiata, 753, 764, 765, 766.
768, 774, 789, 790, 791. 800. 806
Digital processes, 767
Digital tentacles, 767, 769
Dioecism, 740
Diotocardia, 788
Diplomerism, 804, 805
Dorsal fold (of the mantle), 751,
752
Dorsal lobe (of septum), 748
108
828
INDEX.
Efferent branchial vessels (see
Branchio-cardiao vessels)
Eggs, 785, 803, 809
Eledone, 801
Endochondral sinus, 764, 783
Endoskeleton, 766
Enemies (sharks and congers), 739
Enumeration of tentacles, 769
Epidermis, 739, 752
Epipodial Theory, 802
Epipodium, 795, 796, 797, 801, 803,
806
Eschricht, Prof., 800
Excretory products, 790
Experiments, 760
Extrabuccal tentacles, 776, 787
Eye, 781, 793
Faucial follicles, 781
Fiji Islands, 745
Flexure, 797, 798, 799, 804, 806
Food, 774, 808
Foord, A. H., 746
Foot, 739, 795, 798
Fossa buccalis, 767, 776, 787
Fossa infundibularis. 767, 772
Fossa michalis, 767
Fossil Nautilus, 746
Fraisse, P., 793
Free tentacle (of spadix), 777
Function, 760, 768, 796
Funnel, 739, 763, 764, 765, 767,
771, 790, 799, 801, 802, 803, 806
Funnel-cartilage, 766
Funnel organ, 766, 803
Funnel-valve, 765, 803
Gas-production, 747, 760
Gastral ligament, 784
Gastric artei-y, 782
Gastropoda, 739, 753, 766, 789,
791, 795, 796, 802, 804, 806
Generative orifice, 804
Genital arteries, 784
Genital duct, 785
Genital orifices, 754
Genito-intestinal ligament, 757,
760, 784, 806
Geographical distribution, 744
Gill-plates, 788
Gills (see Branchiae)
Gizzard, see Stomach
Glaus (of spadix), 777
Gonad, 784
Gonatus fabricii, 766, 791, 800
Gould, A. A., 743
Grenacher, H., 795, 801, 803
Griffin, L. E., 749, 753, 767, 768,
775, 778, 781, 788
Grobben, C, 801, 803, 803
Growth, 751
Habits, 808
Haemocoel, 759, 780, 806
Hair-eeUs, 778
Haliotis, 753, 788, 789, 792, 794,
795, 796, 802, 806
Haller, B., 755, 757, 758, 759, 764,
784, 787, 790, 798, 807
Earpa, 795, 799, 804
HasweU, W. A., 777
Heart, 755, 805
Hectocotylus, 742
Hedley, C, 745
Hensen, V., 793
Hepatic artery, 782
Historical survey, 736
Homodynamy, 791, 805
Hood, 739, 744, 767, 771, 775
Howes, G. B., 742
Hoyle, W. E., 744, 765, 766, 791,
800
Humboldt, Baron von, 800
Huxley, T. H., 751, 760, 795, 796,
798, 799, 801, 803
Hyatt, A., 748, 760
Hyatt's scar, 748
Hydrostatic apparatus, 747
Hyperpolygyny, 740
Hypobranchial gland, 807
Hypostracum, 7-30
Infrabuecal apparatus, 776, 778, 779
Infi'abuccal nerves, 779
Infrabuecal tentacles, 776
Infundibular fossa, 765 (see also
Fossa iit/undibula7'is)
Infundibular nerves, 766, 779
Infundibular vessels, 764, 783
Initial chamber, 748
Innervation of the ophthalmic
tentacles, 774
Innervation of the osphradia, 792
Intercoelic diaphragm (see Dia-
phragm)
Intereoronal membrane, 776
Intestinal artery, 755
Intra-epidermal blood-spaces, 7.39
Jatta, G., 765, 803
Jhering, H. v., 737, 774, 795, 799,
801, 803, 805
Joubin, L., 745, 752, 781, 789
Keferstein, W., 736, 740, 750, 758,
764, 787, 790
Kerr, J. G., 754, 771, 779, 783,
787, 801, 803, 805
Kerr's collector nerve, 779
KoUiker, A. von, 758
Labial papillae, 767, 776
Labial processes, 767, 776
Labial tentacles (see Extrabuccal
and Infrabuecal tentacles)
Lacaze-Duthiers, H. de, 808
Lankester, 737, 740, 747, 752, 753,
754, 764, 767, 769, 771, 789, 791,
803
Lendenfeld, R. von, 748
Lepidoteuthis grimaldii, 745
Lesser aorta (see Aorta inferior)
Leuckart, K., 801
Levatores infundibuli, 765
Limbus infundibuli. 765
Liver, 780
Locomotion, 739, 790
Loligo, 788, 802, 803
Loven, S., 802
Lymph-glands, 789
Macdonald. J. D., 743, 744, 772,
793, 794
llacrobranchiate segment. 805
Mantle, 746, 751, 753, 790, 803, 805
Marginal appendages, 767, 801
Alargines limitantes, 752
Melanesian Plateau, 745
Membrana basilaris (of the siph-
uncle), 759
Mesopodium, 795, 803
Metamerism (see Diplomerism)
Metapodium, 795, 803
Microbranehiate segment, 804
Milne Bay, 744
Moreton, Hon. M. H., 744
Moseley, H. N., 745, 790, 808
Mucus-gland, 765
Miiller, J., 800
Muscle-lines, 749, 751
Muscle-scar, 749
Nautilus bradfordensis, 743
Nautilus macromphalus, 740, 744,
809
Nautilus pompilius, 740, 743, 777
Nautilus pompilius var. iloretoni,
744
Nautilus stenomphalus, 743, 744
Nautilus umbilicatus, 744, 745, 809
Needhamiau sac, 787
Nephrostomes. 805
Nerve-centres, 764, 768
Nerves of the tentacles, 768, 772
Nervous system, 774
New Caledonia, 745
New Hebrides, 745
INDEX.
829
Nicobar Islands, 743, 745
Nidamental gland, 753, 755, 756,
780, 805, 807, 809
Nuchal membrane, 740, 751, 752,
763, 765, 767, 781
Octopoda, 769, 807
Octopus, 740, 766, 790, 798, 801
Oesophagus, 781
Oken, L., 737
Olfactory nerve, 775
Oliva, 795, 799
Ophthalmic tentacles, 767, 773,
793, 794
Optic ganglion, 764, 765
Oral sheath, 767, 776
d'Orbigny, C, 742, 743
Organ of Owen, 778, 779
Organ of Valenciennes, 771, 779,
787, 809
Organ of Van der Hoeven, 776,
778, 779
Orientation, 798, 806
Oro-anal axis, 797, 805
Osphradia, 753, 756, 790, 804
Ostium ahdominale, 787
Ostracum, 750
Otocones, 794
Otocyst, 744, 764, 765, 772, 781
794
Ovary, 755, 786
Oviduct, 787
Owen, E., 736. 737, 738, 746, 751,
752, 754, 755, 758, 764, 767, 768,
771, 774, 779, 780, 784, 793, 799,
800, 809
Pallial arteries, 754, 762, 784
Pallial complex, 753
Pallial derivatives, 796
Palhal veins, 747, 759, 763
Pallio-nuchal artery, 763, 782
Pallio-septal arteries, 757
Pallio-siphonal sinus, 760
Pallio-viseeral flexure of Nautilus,
806
Pallio-viseeral ligament, 751, 757,
784
Pallio-viseeral region, 804
Pallio-viseeral torsion of Haliotis,
806
Paper Nautilus, 808
Papillae vallatae, 739
Parapodia and Pteropodia, 795
Pearl formation, 746
Pedal derivatives, 796
Pedal ganglion and commissure,
781 (see also Coronal ganglion)
Pedal gland, 766, 803
Pedal Theory, 801
Pelecypoda (see Acephala)
Pelseneer, P., 760, 775, 791, 794,
795, 796, 801, 803
Penis, 754, 787
Pericardial glands (or follicles), 790,
804
Pericardial ligament, 805
Pericardium, 781, 790, 805
Perichondrium, 764
Perineurium, 764
Periostracum, 750
Periotic membrane, 764
Peripheral ganglion cells, 768, 793
Peristomial haemocoel, 765, 766,
780, 783
Perivisceral coelom, 757, 760, 780,
805
Pigment, 739, 752, 768
Pigmentation, 738, 790, 804
Plate, H. L., 792
Platjpodium, 796, 798, 799, 803
Pleural derivatives, 796
Pleural folds, 796
Pleuro-pedal system, 804
Pleurotomaria, 753, 788, 789, 794,
804
Post-anal papilla, 791
Prepuce, 777
Preputial gland, 777
Propagation, 746, 747, 809
Propodium, 795, 803
Prosobranchiata, 753, 796, 804,
806, 807
Protoconch, 748
Protopodium, 795, 801, 803, 806
Proveutrioular artery, 782
Pteropodial lobes, 801, 803 (see also
Parapodia)
Pyrifonn gland, 754, 780, 804
Receptaculiim leminis, 780
Rectum, 780
RecuiTent arteries, 783, 797
Recurrent nerve, 775, 797
Reeve, L., 809
Regeneration, 810
Renal organs, 753, 788, 790, 804
Renal orifices, 753, 754, 804, 807
Reno-branchial vessels (same as
Afferent branchial vessels), 780,
788, 790, 804
Respiration, 790
Rete mirabile, 786
Retractor muscles, 765, 766, 781
Khinophore, 765, 779, 781, 791, 794
Rumphius, G. E., 737, 739
Salivary glands, 781
Scaphopoda, 806
Sepia, 740, 766, 788, 789
Septa of shell, 746, 748
Septal area (of mantle), 750
Septal contour, 749
Septal membrane, 749
Septal suture, 749, 751
Sequence of nerves and tentacles,
772
Sexual dimorphism, 740
Sexual maturity, 748
Shell, 747, 751, 799, 809
Shell-gi-owth, 748
Shell-lines, 749
Shell-muscles, 738, 750
Simroth, H., 806
Sinug venosus', 780, 788
Siphon, 766
Siphonal cartilages, 766
Siphonopodium, 764, 796, 799, 801
Siphuncle, 739, 746, 747, 754, 757,
759, 766, 797, 798
Siphuncular artery, 758
Siphuncular vein, 759, 763
Situs im-ersus, 802, 810
Sowerby, G. B., 743, 808
Spadix, 741, 776, 777, 783, 810
Spengel, J. W., 792
Sperm atophore, 780, 787, 809
Spiruhi, 760, 799, 807
Statistics, 740
Stomach, 755, 784
Stomatogastric nerves, 774, 783
S trombus, 794
Substitution, 766, 796
Suckers of Cephalopoda, 769, 801
Suctorial ridges, 768, 800
SjTiipathetic nervous system, 774
Symphysial tracts, 771, 779, 803
Syncerebrum, 772
Systemic aorta, 763, 781, 784, 805
Teleostean fishes (position of vent),
797
Tentacles, 800 (see also Digital
tentacles)
Tentacular arteries, 783
Tentacular crown, 767
Tessellate tentacle (of spadix), 777
Tetrabranchiata, 764, 807
Thiele, J., 750
Topography, 753, 783, 792, 793,
796, 799, 803
Trochus, 792
Truncus arteriosus, 781
Umbilicus of shell, 743, 748. 809
Valenciennes, A., 736, 748, 750,
830
INDEX.
754, 758, 760, 767, 769, 771, 774,
779, 794, 800
Van der Hoeven, J., 739, 740, 741,
749, 751, 752, 753, 776, 777, 778,
779, 787, 791
Variation, 744, 791, 810
Vas deferens, 787, 810
Vayssi^re, A., 740, 742, 743, 767,
768, 770, 772, 777
Velar Theory, 802
Velum, 801, 803
Vena cavb, 754, 764, 780
Vesicula semitialis, 787
Visceral nerves, 791, 805
Visceral sac, 746, 754, 797, 798
Viscero-brauchial nerves, 792, 804
Viscero-osphradial nerve, 804
Viscero-pericardial orifices, 746,
753, 754, 805
Vitelline membrane, 809
Vitreous body, 794
Vrolik, W., 739, 747, 753. 754.
758
Vulva, 754
Wegmann, H., 753, 788, 789
Whorls of tentacles, 767, 776, 800,
803
WiUey, A., 740, 743. 749, 752, 767,
803
Williams, T., 790
Woodward, B. B., 749
Woodward, M. F., 753, 789, 794, 804
Yolk, 786
Yolk-sac of Cephalopoda, 803
Ziegler, H. E., 805
Zygobranchiata, 753
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